THE  LIBRARY 

OF 
THE  UNIVERSITY 

OF  CALIFORNIA 


PRESENTED  BY 

PROF.  CHARLES  A.  KOFOID  AND 
MRS.  PRUDENCE  W.  KOFOID 


JUNE'192°  MEMOIR  38 


CORNELL  UNIVERSITY 
AGRICULTURAL  EXPERIMENT  STATION 


THE  CRANE-FLIES  OF  NEW  YORK 
PART  II.    BIOLOGY  AND  PHYLOGENY 


ITHACA,  NEW  YORK 
PUBLISHED  BY  THE  UNIVERSITY 


MEMOIR  38 


PLATE  XI 


THEODOR    BELING 

(1816-1898) 


KARL  WILHELM  THEODOR  BELING 

No  sketch  of  the  life  of  Theodor  Beling,  the  great  German  student  of  the 
immature  stages  of  the  Tipuloidea,  has  ever  appeared  in  English.  It  was 
with  considerable  difficulty  that  the  writer  was  able  to  get  into  communication 
with  relatives  and  friends  of  Beling  and  obtain  the  data  presented  herewith.* 

Theodor  Beling  was  born  at  Steterburg,  in  the  Duchy  of  Brunswick, 
Germany,  on  March  26,  1816.  He  was  a  son  of  the  district  forester.  From 
1828  to  1834  he  attended  school  at  Wolfenbiittel.  At  Easter,  1834,  he  began 
his  chosen  career  as  a  forester,  having  served  his  prescribed  apprenticeship 
of  two  years  with  his  father  at  Danndorf.  He  attended  the  Royal  Saxon 
Academy  at  Tharand  from  1836  to  1837,  and  in  the  following  year  visited  the 
University  of  Gottingen  to  round  out  his  technical  training  in  various  branches 
relating  to  forestry.  Beling's  career  as  a  forester  extends  from  Easter,  .1834, 
to  October  1,  1888,  when  he  was  officially  retired  with  a  pension.  In  April, 
1861,  he  was  made  Master  of  the  Forest,  a  position  which  he  held  until  his 
retirement. 

After  his  retirement  from  active  public  service,  Beling  long  continued  his 
vigorous  researches  afield  and  at  home.  When  far  advanced  in  years,  he 
went  alone  on  long  trips  to  seek  new  stations  for  plants  and  to  observe  animal 
life  in  wood  and  field.  He  died  on  December  17,  1898,  at  Seesen,  where 
much  of  his  finest  work  was  accomplished. 

During  his  lifetime  Beling  published  one  hundred  and  seven  articles.  The 
series  of  three  papers  which  rank  him  as  a  pioneer  in  the  field  of  crane-fly 
biology  were  published  in  the  years  1873,  1879,  and  1886,  respectively.  In 
these  papers  the  histories  of  sixty-nine  species  of  crane-flies  are  discussed  in 
detail,  and  brief  notes  on  five  additional  species  are  included.  The  other 
published  articles  of  Beling  cover  a  remarkable  range  of  subjects  relating 
directly  or  indirectly  to  the  science  of  forestry.  His  most  important  researches 
on  the  life  histories  of  insects,  in  addition  to  those  on  the  Tipuloidea,  are  on 
the  coleopterous  families  Parnidae  and  Elateridae.  His  published  articles 
are  dated  between  1850  and  1888,  but  practically  all  of  his  entomological 
studies  were  published  in  the  seventies  and  eighties. 

Beling's  excellent  collection  of  dried  plants,  mounted  birds,  and  sections 
of  various  woods,  and  his  cabinet  of  insects  —  the  last-named  including  the 
types  of  several  species  described  by  himself  and  by  others  —  arc  preserved 
in  the  collection  of  the  Natural  History  Museum  at  Brunswick.  It  is  under- 
stood that  the  insects  in  this  collection,  including  the  alcoholic  larvae  and 
pupae  of  the  Tipuloidea,  are  still  in  excellent  condition. 

*  Sincere  thanks  are  here  extended  to  ths  following  persons,  who  have  added  materially 
to  the  value  of  this  biographical  notice: 

Landesforstmeister  K.  Block,  son-in-law  of  Beling,  who  published  in  April,  1899  (in 
Allgemeinen  Forst-  und  Jagd-Zeitung),  a  complete  obituary  notice  relating  to  his  father- 
in  law,  and  to  whom  the  writer  is  indebted  for  several  additional  unpublished  notes  and 
for  the  loan  of  the  excellent  portrait  of  Peling  reproduced  herewith. 

J.  Meerwartz,  Director  of  the  Natural  History  Museum  at  Brunswick,  who  has  fur- 
nished data  concerning  the  present  condition  of  the  collection  of  the  immature  stages  of 
the  Tipuloidea  described  by  Beling,  now  preserved  in  the  above-mentioned  institution. 

William  Prindle  Alexander  and  Elsa  Miiller  Alexander,  who  have  rendered  valuable 
service  in  translating  the  account  of  Beling's  life. 


142,0 
Bioloqy 

CONTENTS 

General  considerations 704 

Representative  crane-fly  life  histories 704 

Eriocera  longicornis  (Walk.) 704 

Cylindrotoma  splendens  Doane 708 

Life  activities  of  crane-flies 710 

The  adult 710 

The  egg/ 714 

The  larva . . . 715 

The  pupa 719 

Enemies 721 

Predatory  natural  enemies 721 

Vertebrates 721 

Invertebrates ,, .  727 

Parasitic  natural  enemies 730 

External  parasites 730 

Internal  parasites 733 

Historical  summary  of  the  immature  stages 735 

Economic  importance 736 

Species  affecting  herbaceous  plants 737 

Species  affecting  woody  plants 739 

Methods  of  experimental  procedure 740 

Collecting 740 

Killing  and  preserving 740 

Study 741 

Rearing 741 

Types  of  the  immature  stages 743 

External  morphology 744 

The  larva 744 

General  features 744 

Body  form ." 745 

Organs  of  respiration 746 

The  spiracular  disk 746 

Anal  gills 748 

The  head 748 

The  pupa 752 

General  features 752 

The  head 753 

The  thorax 754 

The  abdomen 756 

Internal  morphology 758 

The  digestive  system 758 

Herbivorous  larvae 758 

Carnivorous  larvae 759 

Respiration  and  circulation 760 

The  nervous  system 762 

The  reproductive  system 762 

The  muscular  system 763 

The  fat  bodies,  or  adipose  tissue 763 

The  imaginal  disks,  or  histoblasts 763 

695J 


M34564O 


696  CONTENTS 

PAGE 

Phytogeny 763 

The  geological  record 764 

Comparative  morphology 766 

Phylogenetic  considerations '. 766 

Life  histories  of  crane-flies,  with  keys  and  descriptions  of  the  species 767 

Family  Tanyderidae 769 

Family  Ptychopteridae 772 

Subfamily  Pfcychopterinae 773 

Subfamily  Bittacomorphinae 779 

Family  Rhyphidae 787 

Subfamily  Trichocerinae 789 

Family  Tipulidae 791 

Subfamily  Limnobiinae 793 

Tribe  Limnobiini 795 

Subtribe  Antocharia 799 

Subtribe  Ellipteraria 806 

Subtribe  Limnobaria 808 

Subtribe  Dicranoptycharia 828 

Subtribe  Rhamphidaria 830 

Tribe  Hexatomini 835 

Subtribe  Ularia 838 

Subtribe  Epiphragmaria 843 

Subtribe  Pseudolimnophilaria 848 

Subtribe  Dactylolabaria 852 

Subtribe  Limnophilaria 856 

Subtribe  Hexatomaria 876 

Subtribe  Polymeraria 893 

Tribe  Pediciini 894 

Subtribe  Adelphomyaria 895 

Subtribe  Pedicaria 899 

Tribe  Eriopterini 908 

Subtribe  Eriopteraria 911 

Subtribe  Elephantomyaria 952 

Tribe  Styringomyiini 957 

Subfamily  Cylindrotominae 959 

Subfamily  Tipulinae 974 

Tribe  Tipulini 981 

Subtribe  Dolichopezaria 981 

Subtribe  Ctenophoraria 986 

Subtribe  Tipularia 990 

Bibliography  and  references  cited .- 1019 

.  1129 


THE  CRANE-FLIES  OF  NEW  YORK 
PART  II.     BIOLOGY  AND  PHYLOGENY 


THE  CRANE-FLIES  OF  NEW  YORK 

PART  II.     BIOLOGY  AND  PHYLOGENY1 

CHARLES  PAUL  ALEXANDER 

A  preliminary  classification  of  the  immature  stages  of  the  Tipulidae  and 
related  families,  suggested  to  the  writer  by  Dr.  J.  G.  Needham  in  1911, 
is  presented  in  this  memoir.  But  few  of  the  sixty-odd  families  of  Diptera, 
and  comparatively  few  species  of  the  Tipulidae,  have  as  yet  been  studied 
from  this  viewpoint,  and  therefore  the  arrangement  herein  adopted  must 
be  considered  as  tentative.  The  majority  of  the  specimens  used  in  the 
study  were  reared  in  New  York  State,  the  crane-fly  fauna  of  which  is 
typical  of  a  great  area  thruout  northeastern  North  America.  In  1913, 
thru  the  kindness  of  Dr.  Charles  D.  Woods  and  Dr.  Edith  M.  Patch, 
the  writer  was  enabled  to  continue  his  investigations  in  Maine. 

It  is  the  writer's  purpose  to  outline  the  morphological  characters  avail- 
able for  the  classification  of  the  larvae  and  the  pupae,  and  to  give 
preliminary  keys  for  the  separation  of  the  various  groups;  such  keys  will, 
of  necessity,  require  constant  revision  or  complete  remodeling  with  the 
accession  of  new  life-history  material,  but  it  is  at  least  hoped  that  they 
may  furnish  a  basis  for  future  investigation.  The  most  important  work  on 
the  European  fauna,  that  of  Theodor  Beling  (1873  to  1887) ,2  is  rendered 
incomplete  by  the  total  lack  of  illustration,  the  insufficiency  of  description 
of  the  details  of  the  larval  head,  and  the  artificial  nature  of  the  keys. 
That  this  difficulty  in  using  Beling's  figureless  descriptions  is  not  confined 
to  the  writer  is  shown  by  the  following  criticism  by  one  of  his  fellow- 
countrymen  (Czizek,  1911:7): 

"  Leider  fehlen  uns  bis  jetzt  fast  vollstandig  gute  Abbildungen  der  Larven  und  Puppen, 
ein  fiihlbarer  Mangel  auch  in  Beling's  Abhandlungen,  da  die  genaueste  Beschreibung  das 
Bild  nie  ganz  zu  ersetzen  vermag." 

It  is  intended  to  include  in  this  paper  the  following  material: 
1.  Descriptions  of  all  new  life-history  material  available,  with  notes 
on  the  biology  and  occurrence  of  the  species. 

1  Part  I  of  The  Crane-Flies  of  New  York,  dealing  with  distribution  and  taxonomy  of  the  adult  flies,  was 
published  in  1919  as  Memoir  25  of  the  Cornell  University  Agricultural  Experiment  Station. 

2  Dates  in  parenthesis  refer  to  Bibliography  and  References  Cited,  page  1019. 

699 


700  CHAKLES  PAUL  ALEXANDER 

2.  Brief  summaries  of  published  life  histories  of  genera  and  important 
species  not  available  for  study  as  specimens  and  included  here  to  complete 
the  data. 

3.  Summaries  and  tabulations  of  life-history  records,  larval  habitats, 
economic  importance,  and  related  subjects. 

4.  Keys  to  the  families,  tribes,  and  lesser  groups. 

The  adult  flies  are  not  here  considered  in  any  detail,  since  they  have 
been  discussed  by  the  writer  in  an  earlier  paper  (Alexander,  1919d). 

The  life  histories  remaining  to  be  discovered  in  the  Nearctic  fauna 
are  still  numerous  in  species,  tho  few  in  genera.  There  are  but  four  or 
five  genera  whose  life  histories  when  made  known  may  upset  the  present 
ideas  on  arrangement.  Until  more  is  known  of  these  missing  groups, 
they  must  be  classified  according  to  the  adult  structure. 

It  will  be  noted  that  a  number  of  important  changes  in  nomenclature 
have  been  adopted  in  this  paper.  The  system  hitherto  in  vogue,  based 
entirely  on  the  structure  of  the  imagines,  was  conceived  by  Osten  Sacken 
and  represented  the  culmination  of  research  on  the  structure  and  affinities 
of  the  adult  flies.  A  casual  survey  of  the  immature  stages  is  sufficient 
to  show  the  impossibility  of  many  of  the  groups  hitherto  generally  accepted. 
The  principal  modifications  adopted  in  this  paper  are  as  follows : 

1.  The   erection   of   the   family   Tanyderidae   to   receive    the   genera 
Tanyderus  and  Protoplasa.     These  had  hitherto  been   placed  with  the 
Ptychopteridae,  a  group  to  which  they  are  not  closely  allied. 

2.  The  removal  of  the  genus  Trichocera  from  the  Tipulidae  to  the 
Rhyphidae,  and  the  inclusion  of  the  latter  family  as  one  of  the  four 
existing  families  of  crane-flies. 

3.  In  the  Tipulidae,  the  elimination  of  four  tribes  —  Antochini,  Limno- 
philini,   Dolichopezini,   and   Ctenophorini  —  as   being   based   on   a   con- 
glomeration of  forms  referable  to  other  tribes  or  else  separated  on  an 
insufficient  basis.     The  former  tribe  Antochini  included  members  which 
the    writer  '  now    refers    to    the    Limnobiini     (Antocha,    Rhamphidia, 
Dicranoptycha,  and  other  genera)  and  to  the  Eriopterini   (Teucholabis, 
Elephantomyia) ;    the  Limnophilini   are   too    close    to    the    Hexatomini; 
and  the  tipuline  forms  constitute  a  very  compact  group  which  cannot 
well  be  subdivided  into  tribes. 

4.  The  erection  of  nineteen  sub  tribes,  or  divisions,  to  include  lesser 
groups  of  genera  within  the  tribes.     In  the  following  pages  these  are 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II 


701 


treated  in  what  seems  to  be  their  phylogenetic  sequence  from  the  generalized 
to  the  specialized. 

The  arrangement  of  families,   subfamilies,  and  lesser  groups  may  be 
summarized  as  follows: 


Families 

Subfamilies 

Tribes 

Subtribes 

Representative 
genera 

Eoptychopteridae 
(fossil) 

Eoptychoptera 

Architipula 
f  Tanyderus 
\  Protoplasa 
Ptychoptera 
(  Bittacomorpha 
\  Bittacomorphella 
Rhyphus 
Mycetobia 
Trichocera 
Antocha 
Elliptera 
f  Limnobia 
\  Dicranomyia 
Dicranoptycha 
Rhamphidia 
Ula 
Epiphragma 
Pseudolimnophila 
Dactylolabis 
/  Limnophila 
1  Ulomorpha 
j  Eriocera 
\  Hexatoma 
Polymera 
Adelphomyia 
/  Pedicia 
}  Dicranota 
\  Erioptera 
\  Ormosia 
E  lephantomy  ia 
Styringomyia 
f  Cylindrotoma 
}  Phalacrocera 
\  Dolichopeza 
V  Brachypremna 
\  Ctenophora 
\  Tanyptera 
j  Tipula 
\  Nephrotoma 

Architipulidae 
(fossil)  
Tanyderidae  

Ptychopteridae..  .  . 

I  Bittacomorphinae 

f  Rhyphinae  
<!  Mycetobiinae  .... 
[  Trichocerinae  .... 

Limnobiinae  

Cylindrotominae.  . 
|  Tipulinae  

f  Antocharia  
Ellipteraria  

1  Limnobaria  

|  Dieranoptycharia.  .  . 
1  Rhamphidaria  
'Ularia  
Epiphragmaria  
Pseudolimnophilaria 
Dactylolabaria  

Limnophilaria  
Hexatomaria  

Polymeraria  
(  Adelphomyaria  
1  Pedicaria  

1  Eriopteraria  
{  Elephantomyaria.  .  . 

Tipulidae  

Limnobiini  
Hexatomini.    .  . 

1 
Pediciini 

Eriopterini  
[  Styringomyiini  .  .  . 

Tipulini 

j  Dolichopezaria  
<   Ctenophoraria  
[  Tipularia  

702  CHARLES  PAUL  ALEXANDER 

A  permanent  series  of  the  immature  stages  of  the  species  studied  has 
been  carefully  preserved  in  the  writer's  collection,  while  additional  material 
has  been  placed  in  the  principal  collections  of  the  United  States.  A  series 
of  two  hundred  and  seventy  microscope  slides  showing  details  of  structure 
of  the  larvae  and  the  pupae  has  been  prepared,  representing  some  seventy 
species  arranged  in  forty  genera.  The  typical  larvae  and  pupae 
(nepionotypes  and  neanotypes)  have  been  designated  and  preserved  in 
alcohol  for  future  reference. 

For  the  most  part,  the  life  histories  discussed  in  this  paper  have  not 
been  described  hitherto.  Besides  the  reared  material,  the  writer  has  in 
his  collection  a  very  large  number  of  unknown  larvae  and  pupae,  many 
of  them  representing  interesting  and  un described  types.  In  this  paper 
but  three  or  four  of  these  are  considered,  and  these  only  because  they 
introduce  new  features  of  organization  (Eriopterine  No.  1,  Tipulini  No. 
1  and  No.  2). 

A  vast  amount  of  work  remains  to  be  done  on  the  immature  stages 
of  crane-flies.  Exact,  detailed  life  histories  of  individual  genera  and 
species  are  especially  needed  and  the  reward  for  effort  will  undoubtedly 
be  great.  Nothing  is  known  of  the  tropical  and  the  antipodal  faunas, 
and  their  life  histories  will  probably  be  unraveled  but  slowly,  due  to  the 
inaccessibility  of  most  of  the  regions. 

The  majority  of  specimens  studied  in  the  preparation  of  this  paper 
were  reared  by  the  writer  during  the  past  nine  years.  Many  specimens 
were  received  from  various  sources,  however,  including  many  of  the  most 
desirable  life  histories,  and  these  are  herewith  gratefully  acknowledged: 

Dr.  James  G.  Needham,  Ithaca,  New  York.  The  extensive  collections  received  from  Dr. 
Needham  include  the  material  described  in  the  various  Adirondack  reports,  as  well  as 
numerous  previously  unrecorded  specimens.  In  addition,  the  writer  has  accompanied 
Dr.  Needham  on  many  of  his  field  trips  in  quest  of  the  immature  stages  of  aquatic  insects, 
and  has  received  much  encouragement  and  inspiration  from  this  association. 

Charles  W.  Johnson,  Boston,  Massachusetts.  Immature  stages  of  Aeshnasoma,  Elephant- 
omyia,  Teucholabis,  and  other  species. 

The  late  Frederick  Knab,  Washington,  D.  C.  Immature  stages  of  Rhipidia  bryanti, 
Elephantomyia,  Epiphragma  solatrix,  and  other  species. 

J.  A.  Hyslop,  Hagerstown,  Maryland.     Longurio  (larvae),  Oropeza,  Gnophomyia. 

J.  R.  Malloch,  Urbana,  Illinois.     Rhipidia  bryanti. 

C.  T.  Greene,  Washington,  D.  C.     Protoplasa  (supposition). 

H.  L.  Viereck,  Washington,  D.  C.     Protoplasa  (supposition). 

W.  L.  McAtee,  Washington,  D.  C.  Protoplasa  (supposition);  numerous  larvae  from  the 
Pribilof  Islands. 

William  Lundbeck,  Copenhagen,  Denmark.     Helobia  and  other  species. 

Dr.  C.  Wesenberg-Lund,  Hillerod,  Denmark.  Phalacrocera,  Ptychoptera  paludosa,  Tipula 
maxima,  and  other  species. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  703 

William  G.  Dietz,  Hazleton,  Pennsylvania.     Notes  on  larvae  of  Dicranomyia  macateei  Alex. 

Oskar  A.  Johannsen,  Ithaca,  New  York.     Limnobia  fallax,  and  other  species. 

Carl  Ilg,  Philadelphia,  Pennsylvania.     Notes  on  Rhipidia  fidelis. 

O.  H.  Swezey,  Honolulu,  Hawaii.     Notes  on  Libnotes  and  Styringomyia. 

H.  K.  Munro,  Pretoria,  Transvaal.     Notes  on  Trentepohlia  and  Styringomyia. 

Charles  Bruch,  La  Plata,  Argentina.     Notes  on  Trimicra,  Helobia,  and  Epiphragma. 

D.  B.  Young,  Albany,  New  York.     Notes  on  Gnophomyia. 

J.  T.  Lloyd,  Cincinnati,  Ohio.     Eriocera. 

The  late  C.  Gordon  Hewitt,  Ottawa,  Canada.     The  Canadian-Arctic  Tipulidae,  including 

Tipula  arctica. 

A.  E.  Cameron,  Ottawa,  Canada.     Cylindrotoma  splendens. 
R.  C.  Shannon,   Ithaca,  New  York.     Elephantomyia,  Brachypremna,    Tipula  fuliginosa, 

and  other  species. 

J.  Speed  Rogers,  Grinnell,  Iowa.     Notes  on  Phalacrocera  tipulina. 
Harold  Morrison,  Washington,  D.  C.     Holorusia,  and  Tipula  usitata. 
C.  Hamilton  Kennedy,  Columbus,  Ohio.     Prionocera  and  Rhamphidia. 
J.  Chester  Bradley,  Ithaca,  New  York.     Longurio. 
H.  S.  Barber,  Washington,  D.  C.     Oropeza. 
Adam  Boving,  Washington,  D.  C.    Tanyptera;  notes  on  Helobia. 

In  addition  to  the  preceding,  the  writer  acknowledges  specimens  or 
data  received  from  the  following: 

From  Orono,  Maine:    Edith  M.  Patch,  H.  M.  Parshley,  Cordelia  Stanwood. 

From  Cambridge,  Massachusetts:    Roland  Thaxter. 

From  Ithaca,  New  York:  H.  H.  Knight,  L.  G.  Brown,  C.  C.  Hamilton,  V.  R.  Haber,  E. 
A.  Richmond,  Chih  Ping,  Waro  Nakahara,  M.  D.  Leonard,  W.  A.  Riley,  J.  D.  Tothill, 
Mrs.  J.  D.  Tothill,  W.  P.  Alexander,  W.  A.  Clemens,  W.  C.  Woods,  P.  A.  Munz,  P.  W. 
Claassen,  Axel  Olsson. 

From  Washington,  D.  C.:    C.  H.  Popenoe,  E.  R.  Kalmbach,  Alex.  Wetmore,  A.  T.  Speare. 

From  Lawrence,  Kansas:    H.  B.  Hungerford,  P.  B.  Lawson,  H.  L.  Fackler. 

From  Urbana,  Illinois:    T.  H.  Frison,  W.  P.  Flint. 

The  writer  would  express  his  great  indebtedness  to  Dr.  Ewald  Bergroth, 
of  Jamsa,  Finland,  for  help  in  determining  the  derivation  of  some  of  the 
more  obscure  generic  names  in  the  Tipulidae;  to  Professor  T.  D.  A. 
Cockerell  for  notes  on  the  stratigraphy  of  fossil-bearing  beds  in  many 
parts  of  the  world;  and  to  Miss  Lela  G.  Gross,  assistant  editor  of  the 
Cornell  memoirs,  whose  critical  work  on  this  and  the  preceding  part 
under  this  title  has  been  of  the  greatest  value  in  assuring  uniformity 
and  accuracy. 

Further  acknowledgments  of  assistance  in  determining  plant  and 
animal  associates  of  crane-flies  are  made  thruout  the  text.  In  conclusion, 
the  writer  expresses  his  indebtedness  to  his  wife,  Mabel  M.  Alexander, 
for  her  untiring  care  and  assistance  in  the  typing  and  final  preparation 
of  this  thesis. 


704  CHARLES  PAUL  ALEXANDER 

GENERAL  CONSIDERATIONS 
REPRESENTATIVE    CRANE-FLY   LIFE   HISTORIES 

The  life  histories  of  but  few  species  of  crane-flies  have  been  studied 
in  detail.  The  very  nature  of  the  habitat  (mud  or  earth)  required  by 
most  species  of  the  family  renders  it  a  most  difficult  operation  to  rear 
the  species  from  the  egg  to  the  adult  and  note  the  various  stages,  their 
molts,  their  habits,  and  other  features.  The  subfamily  Cylindrotominae, 
the  immature  stages  of  which  live  on  the  leaves  of  various  higher  plants 
and  curiously  resemble  the  caterpillars  of  certain  Lepidoptera,  furnishes 
species  whose  habits  are  more  readily  studied  than  most  others,  and  as 
a  result  the  immature  stages  of  this  group  are  better  known,  perhaps, 
than  those  of  any  other  section  of  the  family.  Two  widely  different 
species  have  been  chosen,  and  their  life  histories  as  they  are  known  at 
present  are  here  outlined.  The  first  is  a  species  of  Eriocera,  a  powerful, 
semi-aquatic  carnivore;  the  second  is  a  species  of  Cylindrotoma,  a  ter- 
restrial herbivore.  In  the  text  which  follows,  notes  on  the  life  activities 
of  various  other  species  are  given,  but  the  gaps  in  the  knowledge  of  this 
phase  of  the  subject  are  very  considerable  and  there  still  remain  innumer- 
able interesting  facts  to  be  ascertained. 

Eriocera  longicornis  (Walk.) 

The  common  crane-fly  Eriocera  longicornis  is  widely  distributed  over 
the  northeastern  United  States  and  Canada.  Altho  the  species  is  local 
in  its  distribution,  the  flies  may  be  found  in  abundance  wherever  it  does 
occur.  The  situations  that  favor  the  presence  of  these  flies  are  large 
streams  or  rivers  with  sand  or  gravel  bottoms  and  banks.  The  following 
notes  were  taken,  partly  in  Fulton  County,  New  York,  along  the  Sacan- 
daga  River,  and  partly  in  Tompkins  County,  along  Fall  and  Cascadilla 
Creeks.  Some  of  these  data  have  already  been  published  by  Alexander 
and  Lloyd  (1914:12-18)  and  by  the  writer  (Alexander,  1915  c:  149-152). 

On  May  27,  1914,  the  adult  flies  were  exceedingly  numerous  near  the 
village  of  Northampton,  Fulton  County.  They  were  present  in  untold 
myriads,  and  at  every  step  they  arose  in  clouds  from  under  foot  or  from 
the  leaves  of  chokecherry  on  which  they  rested.  They  sat  on  the  leaves 
with  the  head  directed  away  from  the  observer,  ready  to  take  instant 
flight,  and  at  the  first  approach  of  a  possible  enemy  they  darted  up  into 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  705 

the  air  and  far  overhead.  A  few  were  in  copulation  on  the  leaves,  but 
from  observations  made  later  it  seems  probable  that  mating  begins  in 
the  air  and  the  united  pair  seek  a  support  later  on.  Toward  twilight 
the  flies  may  be  found  in  great  numbers  in  company  with  many  kinds  of 
caddice  flies  and  may  flies.  The  swarms  vary  in  numbers  from  about 
fifty  individuals  to.  those  including  many  thousands.  Some  of  the  larger 
of  these  swarms  cover  a  vertical  height  of  at  least  fifty  feet,  the  lower 
individuals  being  about  fifteen  feet  above  the  water.  When  danger 
approaches,  the  swarm  either  mounts  into  the  air  overhead  or  retreats 
before  the  breeze,  never  advancing  nor  moving  sidewise. 

Out  over  the  land  in  the  smaller  swarms,  copulation  was  observed 
several  times.  The  males  in  the  swarm  dart  swiftly  at  the  females  and 
seize  them  almost  instantly.  They  then  usually  leave  the  swarm  and  go 
sailing  away,  the  male  above  doing  the  flying,  the  smaller  female  hanging 
limply  beneath.  The  antennae  of  the  male  at  this  time  are  directed 
straight  ahead  and  are  slightly  divergent,  the  legs  hang  downward,  the 
wings  vibrate  rapidly.  The  female  hangs  downward  with  the  legs  trailing 
limply  beneath,  the  wings  nearly  horizontal  and  motionless.  The  dead 
weight  of  the  female  continually  pulls  the  male  toward  the  water,  and 
often  both  fall  into  the  river.  As  a  rule',  copulation  ends  before  the  male 
is  exhausted.  The  male  opens  his  forceps  and  the  female  drops  straight 
downward  for  a  foot  or  so,  quite  like  a  parachute  released  from  a  balloon. 
If  the  pair  are  near  the  surface  at  the  time  of  separation,  the  released 
female  drops  into  the  water.  The  male  darts  upward  again  and  back 
into  the  swarm.  The  female  slowly  flies  away,  usually  upstream  but 
sometimes  downstream,  presumably  to  lay  her  eggs.  She  does  not  stop 
for  swarms  that  she  may  encounter,  and  may  even  make  a  wide  detour 
in  order  to  avoid  them.  In  a  very  few  cases  the  female  is  the  active 
partner  and  succeeds  in  pulling  the  male  where  she  wills  altho  she  is  much 
smaller.  No  matings  were  observed  during  the  hours  of  sunlight,  and 
it  was  only  at  twilight  or  just  after  sunset  that  mating  took  place.  Often 
a  second  male  will  seize  a  female  already  in  copula  and  the  three  will 
come  tumbling  down  into  the  water  together.  One  such  pairing  of  three 
individuals  came  down,  and  when  they  were  about  a  foot  above  a  board 
in  the  water  the  male  in  copulation  dropped  the  other  two.  These  rested 
for  a  moment,  and  then  the  remaining  male  attempted  to  engage  the 
female  in  copulation.  She  resisted  but  finally  he  managed  to  seize  her 


706  CHARLES  PAUL  ALEXANDER 

with  his  forceps.  Then  he  attempted  to  fly  away,  but  she  seized  hold 
of  the  board  with  all  her  feet  and  he  was  unable  to  disengage  her.  This 
seems  to  indicate  that  the  normal  place  for  copulation  is  in  the  air. 

The  motions  of  the  insects  in  the  swarm  were  very  rapid,  almost  like 
those  of  bees,  and  the  sound  produced  was  at  a  very  low  pitch,  much 
lower  than  that  made  by  Culex.  The  movements  are  on  a  horizontal 
plane,  each  individual  flying  mostly  in  the  path  of  a  figure  8,  sometimes 
slowly  and  at  other  times  much  more  rapidly. 

Many  specimens  were  seen  dipping  down  into  the  water,  as  tho  engaged 
in  laying  eggs.  All  of  the  few  specimens  captured  proved  to  be  males, 
but  why  this  sex  should  go  thru  these  motions  is  not  clear  to  the  writer. 
This  action  has  been  observed  several  times  in  various  species  of  crane- 
flies.  It  is  very  probable  that  the  female  lays  her  eggs  in  the  water  in 
this  manner. 

The  eggs  are  pale  white  or  brown,  not  heavily  chitinized  as  are  those 
of  Hexatoma  but  with  the  chorion  feebly  sculptured.  They  vary  in 
number  from  892  to  1034,  with  an  average  of  952.  They  are  small,  about 
the  same  size  as  those  of  Hexatoma.  The  ovaries  almost  completely 
fill  the  abdominal  cavity,  and  the  eggs  are  arranged  in  the  ovaries  like 
bananas  on  a  stalk,  with  numerous  pale  nurse-cells  in  between. 

The  larval  life  is  passed  in  streams,  usually  under  rocks.  The  winter 
is  spent  in  the  larval  condition,  but  the  larvae  do  not  attain  full  size  until 
the  following  spring.  At  this  time  they  come  to  the  land  and  live  in 
the  sand  and  gravel  along  the  banks  of  the  streams.  By  the  alternate 
extension  and  contraction  of  the  body  and  the  inflation  of  the  penultimate 
segment  of  the  abdomen  at  the  moment  of  extension,  the  larvae  are 
capable  of  inflating  this  segment  into  an  enormous  globular  structure 
which  serves  as  an  aid  to  progression  thru  the  soil.  The  food  of  the  larvae 
consists  largely  of  animal  matter,  and  often  large  species,  such  as  chiro- 
nomid  larvae,  are  swallowed  whole.  The  almost  total  lack  of  chitinization 
of  the  mental  region  allows  for  great  distension  of  this  part  of  the  body. 
The  powerful  mandibles  and  the  retrorsely  roughened  esophagus  serve 
the  function  of  both  holding  the  prey  and  preventing  its  ejection  when 
once  swallowed.  Considerable  gravel  and  particles  of  vegetable  tissue 
are  also  found  in  the  pro  ventricular  region. 

When  ready  to  transform  to  the  pupal  condition,  the  larva  becomes 
sluggish.  After  molting  the  last  larval  skin,  the  pupa  is  disclosed,  pale 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  707 

yellowish  white  and  very  callow.  The  pupa  forms  burrows  in  the  loose 
gravel,  these  being  vertical,  or,  more  often,  a  little  oblique.  The  diameter  of 
the  burrow  is  a  little  greater  than  that  of  the  pupa.  The  length  varies,  the 
burrow  for  a  young  pupa  being  a  mere  chamber  inclosing  the  individual 
and  located  from  one-half  to  one  inch  below  the  surface.  As  the  pupa 
becomes  older,  by  a  bobbing  up-and-down  motion  it  lengthens  the  burrow 
upward  until  finally  it  penetrates  the  surface  layer  and  forms  a  small 
shot-like  opening.  Here  the  pupa  rests,  often  bobbing  up  and  down 
with  a  rhythmic  motion,  but  ducking  down  into  the  burrow  when  danger 
threatens.  The  length  of  the  burrow  is  rarely  more,  but  usually  less, 
than  twice  the  length  of  the  pupa.  A  few  of  the  pupae  seem  to  be 
inclosed  in  a  very  delicate  silken  tube  and  the  salivary  glands  seem  to 
be  well  adapted  for  the  purpose  of  forming  silk;  but  the  great  majority 
of  the  pupae  are  entirely  naked.  The  indoor  pupal  period  varies  from 
one  hundred  and  seventy  to  one  hundred  and  seventy-three  hours,  or  a 
little  more  than  seven  days.  The  outdoor  pupal  period,  however,  is 
undoubtedly  longer,  unless  the  weather  is  very  warm. 

The  emergence  of  the  adults  usually  takes  place  during  the  late  hours 
of  the  morning,  the  greatest  number  emerging  between  ten  o'clock  and 
noon.  When  ready  to  emerge,  the  pupa  pushes  part  of  its  body  out  of  the 
earth,  the  posterior  two-thirds  or  half  remaining  attached  to  the  soil. 
If  it  projects  farther  than  this,  its  transformation  seems  to  be  a  very 
difficult  operation.  The  pupa  bends  backward  and  forward  constantly, 
flexing  the  body  dorso-ventrally.  This  motion  appears  to  exhaust  it, 
since  it  frequently  rests.  The  skin  splits  lengthwise  up  the  mesonotum 
and  the  adult  emerges.  The  male  has  difficulty  in  extricating  its  very 
long  antennae  from  their  sheaths.  The  tips  of  the  fore  femora  are  placed 
underneath  the  sharp  spines  of  the  flagellum,  aftd  by  raising  the  legs  the 
insect  pulls  the  antennae  slightly  outward.  These  spines  are  regularly 
spaced,  and,  since  both  fore  legs  work  in  unison,  the  spines  function  as  cogs 
and  the  whole  antenna  is  gradually  forced  from  the  pupal  sheath.  The  body 
is  ca-rried  very  straight  and  stiff  during  the  operation,  and  the  abdomen 
is  very  long  and  pale.  The  drawing  out  of  the  extreme  tips  of  the  antennae 
is  usually  accomplished  by  the  insect  flexing  its  whole  body  backward. 
When  the  antennae  are  freed,  the  insect  walks  a  few  steps  from  the  cast  skin, 
withdrawing  its  abdomen  from  the  case.  (The  emergence  of  E.  spinosa 
is  shown  in  Plate  XII,  1.)  A  drop  of  nearly  colorless  liquid  is  excreted 


708  CHARLES  PAUL  ALEXANDER 

from  the  body  at  this  time.  The  teneral  adult  then  waits  quietly  until  it 
gains  more  strength  and  color.  This  condition  of  the  insect  is  the  most 
dangerous  period  of  its  existence,  since  it  is  defenseless  against  all  enemies. 

The  insects  are  associated  in  the  gravel  with  ground  beetles  of  the 
genera  Omophron,  Schizogenius,  Dyschirius,  Bembidion,  Tachistodes; 
with  click  beetles  of  the  genus  Cryptohypnus;  and  with  rove  beetles,  of 
which  Paederus,  Lathrobium,  and  Gastrolobium  are  the  commonest  forms. 
In  addition,  numbers  of  larvae  of  Tabanidae,  Leptidae  (Atherix),  Eriocera 
spinosa,  E.  fultonensis,  E.  cinerea,  and  Erioptera  armata  were  found. 
Natural  enemies  of  the  pupae  and  the  teneral  imagines  are  the  medium- 
sized  black  lycosid  spiders,  which  preyed  in  numbers  on  the  weak, 
uncolored  adults.  Dozens  of  these  spiders  were  noticed  with  individuals 
of  the  crane-flies  in  their  grasp.  When  alarmed  they  would  run  rapidly 
away,  but  only  in  exceptional  cases  would  they  release  their  victims.  A 
few  spiders  of  other  families,  notably  the  Attidae,  were  found  with  Erio- 
cerae.  Dragon-flies  appear  to  be  the  most  serious  enemies  of  the  active 
adults.  Helocordulia  uhleri  (Selys)  has  been  observed  capturing  the  crane- 
flies  by  darting  back  and  forth  thru  the  swarms  of  individuals. 

Cylindrotoma  splendens  Doane 

The  life  history  of  the  species  Cylindrotoma  splendens  has  been 
worked  out  in  considerable  detail  by  Dr.  A.  E.  Cameron,  from  material 
obtained  near  Westholme,  Vancouver  Island,  British  Columbia,  in  late 
April,  1917.  The  following  account  is  abstracted  from  Cameron's  detailed 
paper  (1918)  on  this  interesting  crane-fly: 

The  adults  first  appear  on  the  wing  about  the  middle  of  May.  Without  food  they  do 
not  live  longer  than  five  or  six  days,  but  in  breeding  cages,  where  they  were  supplied  with 
food  in  the  nature  of  a  sugar  solution,  they  lived  as  long  as  from  seven  to  nine  days.  In 
nature  the  adults  were  found  on  the  wing  during  a  period  of  about  three  weeks. 

Soon  after  emergence  the  adults  begin  to  copulate,  and  one  male  may  have  intercourse 
with  more  than  one  female.  Copulation  often  takes  place  in  a  vertical  position,  the  female 
above,  the  male  below,  with  the  tips  of  the  abdomens  interlocked.  At  times  the  female 
was  noted  hanging  to  the  roof  of  the  breeding  cages,  with  the  male  suspended  head  down- 
ward, his  body  at  an  angle  of  90°  to  that  of  the  female,  and  his  legs  unsupported.  If  dis- 
turbed, the  female  may  walk  off,  dragging  her  mate  after  her,  or  she  may  take  flight,  bearing 
the  male  with  her.  In  nature  the  act  of  copulation  is  generally  undertaken  in  the  deep 
shade  of  the  large  leaves  of  the  insect's  food  plants,  the  sexes  resting  on  the  under  surface 
of  the  leaves  or  on  the  stems.  Copulation  may  last  but  a  few  minutes  or  may  require 
several  hours. 

The  eggs  are  of  a  dull  glistening  white,  elongate-oval  in  shape.  Under  natural  conditions 
they  are  almost  invariably  found  on  the  under  surface  of  the  leaves  of  the  food  plant 
Trautvetteria  grandis  Nuttall  (Ranunculaceae) ,  inserted  beneath  the  incised  epidermis.  They 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  709 

are  usually  deposited  in  series  along  and  just  within  the  margin  of  the  serrate,  palmately- 
lobed  leaf.  They  are  found  in  groups  of  one  or  more,  all  arranged  parallel  to  one  another 
and  with  their  long  axes  perpendicular,  or  nearly  so,  to  the  margin  of  the  leaf.  The  eggs 
are  only  partly  hidden  beneath  the  epidermis,  being  exposed  dorsally,  the  margins  of  the 
slit  overlapping  the  egg  laterally  and,  to  a  lesser  degree,  both  anteriorly  and  posteriorly. 
This  arrangement  of  the  eggs  in  series  gives  to  the  leaf  margin  a  somewhat  beaded  appearance. 
When  the  eggs  are  older,  the  leaf  margin  turns  brown  and  the  presence  of  the  eggs  is  readily 
detected.  In  ovipositing,  the  female  rests  on  the  under  side  of  the  leaf,  with  the  tip  of  the 
abdomen  directed  toward  the  leaf  edge.  The  abdomen  is  slightly  flexed  ventrally,  and  the 
margin  of  the  leaf  is  held  between  the  bifurcated  valves  of  the  ovipositor,  which  is  applied 
to  the  upper  surface  of  the  leaf,  while  the  paired  cutting  valves,  with  their  blades,  are  applied 
against  the  under  surface.  These  blades  are  then  moved  to  and  fro,  cutting  a  slit  in  the 
epidermis.  The  eggs  are  then  deposited.  In  no  case  do  the  eggs  actually  touch  one  another 
on  the  leaf.  In  the  breeding  cages  females  lay  their  eggs  indiscriminately  on  both  the  upper 
and  the  lower  surface  of  the  leaves.  In  some  cases,  when  a  leaf  has  been  eaten  earlier  in 
the  season  by  larvae,  the  female  will  deposit  her  eggs  along  the  ragged  edge  of  this  damaged 
part. 

The  indoor  duration  of  the  egg  stage  is  from  fourteen  to  eighteen  days,  but  in  the  field 
as  long  a  period  as  three  weeks  may  be  required.  When  first  deposited  the  egg  is  translucent, 
grayish  white,  and  spindle-shaped,  with  the  chorion  unornamented,  and  measures  on  an 
average  0.84  by  0.303  millimeter. 

The  larva  requires  about  three  hours  to  emerge  from  the  egg.  The  chorion  of  the  egg 
splits  longitudinally  down  the  mid-dorsal  line,  the  slit  extending  almost  half  the  length  of 
the  egg.  In  emerging,  the  almost  transparent,  grayish  white  larva  utilizes  the  body  tubercles 
as  levers  in  freeing  itself  from  the  eggshell.  As  soon  as  it  is  freed  from  the  shell,  the  newly 
hatched  larva  begins  to  feed  on  the  leaf  tissue  of  the  host  plant. 

The  first-stage  larvae  are  grayish  white  in  color,  are  translucent,  and  measure  1.19  by 
0.37  millimeter  in  size.  They  feed  on  both  the  upper  and  the  lower  surface  of  the  leaf, 
burrowing  thru  the  epidermal  layer  with  their  mandibles  and  feeding  on  the  parenchymatous 
tissue  within.  The  young  larvae  are  very  sluggish  and  are  not  readily  disturbed  when 
feeding,  the  mandibles  being  firmly  embedded  in  the  leaf  tissue.  While  engaged  in  feeding 
they  assume  various  positions  on  the  leaf  surface.  At  the  end  of  nine  days  the  larvae  have 
increased  in  length  to  5.84  millimeters,  and  show  all  the  characteristic  behavior  of  the  full- 
grown  larvae.  Growth  is  very  slow,  and  before  the  first  molt  the  larvae  become  covered 
with  particles  of  their  excrement,  which  adheres  readily  to  the  skin.  The  first  larval  molt 
occurs  after  a  period  of  from  eighteen  to  twenty-one  clays;  in  some  cases,  however,  it  does 
not  occur  until  five  or  six  weeks  have  elapsed. 

The  second-stage  larvae  gradually  assume  a  leaf-green  color  as  they  continue  to  feed. 
Toward  the  end  of  July,  coincident  with  the  dying-off  of  their  food  plant,  the  larvae,  which 
have  now  reached  a  length  of  from  8.32  to  9  millimeters,  become  quiescent  and  cease  to 
feed.  Feeding  and  movement  gradually  cease  completely  and  the  larvae  remain  clinging 
motionless  to  the  leaves.  As  the  leaves  wither,  the  larvae  drop  off,  and,  in  some  cases, 
attach  themselves  to  the  stems;  under  natural  conditions,  however,  they  usually  fall  among 
the  dead  leaves  on  the  ground  and  under  these  they  pass  the  whiter  in  a  dormant  condition. 
Many  of  these  larvae  match  the  brown  color  of  the  dead  leaves,  but  some  of  them  retain 
their  leaf-green  tint. 

The  overwintering  larvae  first  show  signs  of  activity  in  March  of  the  following  spring, 
when  Trautvetteria  grandis  sends  up  its  new  shoots.  Growth  then  proceeds  rapidly  until 
pupation  in  the  middle  of  May.  There  seems  to  be  one  molt  before  hibernation  and  two 
after,  the  last  being  the  casting  of  the  larval  skin,  previous  to  pupation.  The  fully  grown 
larvae  measure  17  millimeters.  They  are  invariably  found  on  the  upper  surface  of  the  leaf, 
and  in  the  spring  are  actively  engaged  in  feeding.  On  a  fresh  leaf  the  larvae  usually  begin 
by  skeletonizing  it,  leaving  the  lower  epidermis  intact.  Later  on,  however,  large  holes  may 
be  eaten  completely  thru  the  leaf.  The  fully  grown  larvae,  and  to  a  somewhat  lesser  degree 
the  younger  larvae,  progress  by  a  looping  motion,  which  may  be  aptly  compared  to  that 


710  CHAELES  PAUL  ALEXANDER 

of  the  measuring  worms  (Geometridae) ,  progression  being  accomplished  by  the  aid  of  the 
mandibles  and  the  ventral  tubercles,  or  pseudopodia.  In  a  quiescent  condition  the  thoracic 
region  of  the  body  has  a  noticeable  humped  appearance.  The  fully  grown  larvae  are  very 
sluggish  and  inactive.  When  disturbed  they  relax  their  hold  on  the  leaf  surface  and  fall 
to  the  ground.  Altho  several  hundred  adults  were  reared,  not  a  single  parasite  was  discovered. 

Before  pupation  the  larva  attaches  itself  firmly  to  the  surface  of  the  leai  or  to  the  le£f 
petiole,  by  means  of  its  anal  pseudopodia.  The  skin  splits  transversely  behind  but  is  only 
partially  sloughed  off.  The  head,  the  thorax,  and  the  first  four  abdominal  segments  of  the 
pupa  are  exposed,  but  the  apex  of  the  abdomen  remains  encased  in  the  larval  skin,  the 
terminal  part  of  which,  collapsed  and  wrinkled,  is  attached  to  the  leaf  surface.  Pupation 
may  take  place  on  either  the  upper  or  the  lower  surface,  but  it  occurs  oftener  on  the  former. 
If  pupation  takes  place  on  the  petiole,  it  is  generally  at  the  axil.  In  the  breeding  cages  the 
duration  of  the  pupal  period  was  found  to  vary  from  six  to  ten  days. 

When  the  adult  first  emerges  it  is  of  a  pale  green  color,  which  is  gradually  replaced  by  the 
black  and  yellow  of  the  fully  colored  insect.  From  one  and  one-half  to  two  hours  are  required 
for  the  adult  to  emerge.  After  emergence  is  completed,  the  fly  rests  for  a  short  period 
until  the  cuticle  hardens  and  the  wings  expand.  It  seems  that  the  adherence  of  the  larval 
skin  to  the  pupa  is  necessary  for  the  emergence  of  the  adult,  at  least  in  many  cases.  There 
appears  to  be  a  large  disproportion  of  females  over  males,  this  sometimes  being  as  high  as 
five  to  one.  Since  a  single  male  may  copulate  with  several  females,  this  disproportion  is 
not  so  serious  as  it  appears  at  first  sight. 

LIFE   ACTIVITIES   OF    CRANE-FLIES 

The  adult 

Emergence. —  Emergence  from  the  pupal  hull  may  require  but  a  few 
seconds  (as  described  for  GnophomyiabyHyslop  in  litt.,  the  whole  operation 
requiring  but  eight  seconds),  or  it  may  take  several  hours.  The  emergence 
of  the  strictly  aquatic  genus  Antocha  has  not  been  observed,  but  it  must  be 
practically  instantaneous  as  in  Blepharocera  and  the  lotic  caddice  flies. 

Mating. —  In  several  widely  different  species,  the  females  as  they  emerge 
from  the  pupal  hulls  are  at  once  seized  in  copulation  by  the  males  altho 
they  are  still  callow  and  uncolored.  Mik  (1882  b:  40,  and  1886  a)  discusses 
this  curious  condition  in  considerable  detail.  In  all  the  cases  that  are 
known  to  the  writer  —  Dicranomyia  trinotata,  Discobola  caesarea,  and 
Cylindrotoma  distinctissima  (Mik,  1886  a,  the  last-named  also  cited  by  Mik, 
1882  b),  Liogma  glabrata  (Miiggenberg,  1901),  Tipula  rufrna  (Giard,  1895), 
and  Tipula  ultima  (Caudell,  1913)  —  the  abdomen  of  the  female  is 
elongate,  flabby,  and  nearly  colorless.  In  Dicranomyia  trinotata  the 
females  scarcely  have  time  to  remove  their  legs  from  the  pupal  sheaths 
before  they  are  seized  in  copulation.  In  other  cases  the  males  emerge 
before  the  females  and  wait  beside  the  pupae  for  the  emergence  of 
their  mates,  when  they  at  once  seize  them  in  copula.  In  most  species 
of  Tipula  the  males,  when  seeking  the  females,  progress  by  a  fluttering 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  711 

motion,  partly  flying  and  partly  walking,  over  the  ground  or  up  the  trunks 
of  trees.  This  habit  is  discussed  under  the  account  of  Tipula  taughannock 
(p.*1013),  and  has  been  observed  in  other  woodland-inhabiting  species  of 
this  genus  —  as  T.  macrolabis,  T.  fuliginosa,  T.  fragilis,  and  others.  In 
T.  fragilis,  when  a  male  comes  upon  a  pair  already  in  copula  he  passes 
on  without  interrupting  them.  Somewhat  similar  mating  habits  are  found 
in  some  species  of  Dicranomyia  (D.  trinotata,  D.  badia,  and  D.  simulans), 
Discobola,  Antocha,  Chionea,  Dactylolabis  montana,  some  Pediciini  as 
Dicranota,  and  the  Cylindrotominae. 

Many  crane-flies  have  developed  swarming  habits  for  the  purpose  of 
mating,  these  including  representatives  of  most  of  the  tribes  of  the 
Limnobiinae  and  a  few  tipuline  forms.  Dicranomyia  morioides  was 
observed  by  Needham  (1908 a: 204)  swarming  in  vast  numbers  near 
Ithaca,  New  York,  but  here  the  swarms  consisted  only  of  males.  Like- 
wise, Erioptera  armata  (Needham,  1908 a: 206)  was  found  swarming  near 
Lake  Forest,  Illinois;  but,  out  of  several  hundred  individuals  captured, 
all  except  three  were  males.  The  writer  has  observed  swarming  in  numer- 
ous species  of  Ormosia,  Molophilus,  Erioptera,  Gonomyia,  Rhabdomastix, 
Limnophila,  Ula,  Epiphragma,  Eriocera,  Dicranota,  Rhaphidolabis, 
Trichocera,  and  other  genera,  and  here,  too,  the  males  were  always 
predominant.  The  specific  data  may  be  consulted  under  these  various 
headings.  The  males  of  Dicranota  swarm  in  rather  large  numbers  pre- 
liminary to  searching  for  the  females,  which  rest  quietly  on  the  branches 
of  neighboring  shrubbery.  Limnophila  ultima,  as  noted  at  Gloversville, 
New  York,  on  September  7,  1916,  was  swarming  at  half  past  six  o'clock 
in  the  evening.  The  swarms  here  consisted  of  from  fifty  to  sixty  individuals 
and  took  place  from  ten  to  eighteen  feet  above  the  earth.  Mating  took 
place  frequently  in  the  air,  and  as  soon  as  a  pair  were  in  copula  they  flew 
away  to  some  point  to  rest,  many  pairs  being  observed  hanging  on  a 
clothesline  a  few  feet  away.  There  were  three  distinct  swarms,  which 
showed  little  tendency  to  fuse  altho  their  flight  area  was  very  close.  The 
vast  swarms  of  Eriocera  longicornis  and  of  Trichocera  are  mentioned  or 
discussed  elsewhere  in  this  paper.  When  pairs  are  in  copula,  they  readily 
take  flight,  still  united,  the  female  usually  trailing  the  male  after  her; 
altho  in  a  few  groups,  in  which  the  male  is  the  larger  individual  of  the 
two,  the  situation  is  the  opposite.  Brachypremna,  the  familiar  "  weaver  " 
of  the  Southern  States,  has  a  very  remarkable  vertical  dance  of  several 


712  CHARLES  PAUL  ALEXANDER 

feet  in  shady  spots,  and  has  been  aptly  termed  by  Johnson  (1907-12 
[1909] :  123)  "  the  king  of  the  dancing  tipulids."  Doubtless  many  interest- 
ing facts  remain  to  be  discovered  concerning  the  dances  of  the  tropical 
species  of  Brachypremna,  Tanypremna,  and  Megistocera. 

Dancing. —  The  dances  of  Thrypticomyia  saltens  (Dol.)  should  be 
mentioned  at  this  point.  According  to  Doleschall  (1857),  Jacobson 
(De  Meijere,  1911:22-23),  and  others,  this  species  is  common  in  Java 
in  shady  places  thruout  the  year.  The  insects  have  the  habit  of  clinging 
to  spider  webs  by  means  of  the  fore  feet,  or,  if  a  fore  foot  is  lacking,  one 
of  the  middle  legs  is  used.  Often  twenty  or  more  of  these  flies  are  seen 
hanging  close  beside  one  another  on  a  horizontally  spun  web,  all  seesawing 
rapidly  up  and  down  and  at  the  same  time  swaying  to  and  fro,  sometimes 
rhythmically,  sometimes  not.  This  ludicrous  tight-rope  dance  is  con- 
tinued for  a  long  time.  Somewhat  similar  habits  have  been  described 
for  Trentepohlia  pennipes,  Rhamphidia  venustissima  Alex., and  other  species 
with  white  tarsi. 

Bobbing. —  Species  of  Dicranomyia  and  Geranomyia,  as  well  as  a  few 
other  crane-flies,  have  the  curious  habit  of  bobbing  up  and  down  while 
resting,  the  long,  slender  legs  acting  as  springs.  The  species  of  the  latter 
genus  practice  this  same  oscillating  movement  while  resting  on  a  head 
of  flowers  and  feeding.  H.  K.  Munro  has  recorded  this  bobbing  habit 
in  a  species  of  Trentepohlia  (p.  943). 

Resting  habits. —  When  at  rest  crane-flies  assume  various  positions 
which  are  often  fairly  characteristic.  Many  species  (Erioptera,  Molo- 
philus,  and  some  Dicranomyia)  resemble  spiders  when  flattened  against 
a  tree  trunk  or  some  other  vertical  support.  Styringomyia  resembles 
a  bit  of  cobweb,  the  fore  and  middle  legs  stretched  out  in  front,  the  hind 
legs  directed  backward.  In  a  position  of  rest  the  tipuline  forms  generally 
hold  their  wings  outspread  or  divaricate,  exceptions  being  in  the  genera 
Longurio,  Oropeza,  and  others,  and  in  a  few  species  of  Tipula,  as  T.  arctica 
and  the  woodland-inhabiting  species  of  the  marmorata  group  (T.  fragilis 
and  T.  ignobilis) .  The  limnobiine  forms  usually  fold  the  wings  incumbent 
over  the  abdomen,  but  here  again  exceptions  are  found  in  Pedicia, 
Limnophila  toxoneura,  and  other  species,  which  normally  rest  with  the 
wings  outspread.  Oropeza  hangs  to  the  roofs  of  bridges,  culverts,  and 
similar  places,  with  only  the  fore  legs  attached  to  the  support,  the  middle 
legs  divergent,  the  posterior  legs  hanging  loosely  behind.  The  apparently 


THE  CRANE-FLIES  OF  NEW  YORK  — PART  II  713 

closely  allied  Dolichopeza,  on  the  contrary,  has  the  four  anterior  legs 
on  the  support,  the  hind  legs  dangling,  the  wings  divaricate.  Many  of 
these  species  —  as  Oropeza,  Dolichopeza,  Dicranomyia  badia,  and  exotic 
species  of  the  genera  Thrypticomyia  and  Trentepohlia  —  habitually  rest 
on  spider  webs  (page  982;  also,  Knab,  1912).  During  heavy  rains,  crane- 
flies  rest  on  the  lower  side  of  the  broad  leaves  of  deciduous  trees  or  hide 
Deneath  loose  flakes  of  bark. 

Feeding.— The  adult  flies  feed  but  little,  the  majority  of  the  species 
whose  habits  are  at  all  known  merely  lapping  nectar  from  open  flowers. 
Knuth  (1909:579)  and  Wahlgren  (1917)  record  the  plants  frequented  by 
a  number  of  European  Tipulinae  as  well  as  by  Ptychoptera.  These  species 
are  found  on  Umbelliferae  (Aegopodium,  Anthriscus,  Heracleum,  Carum, 
Anethum,  and  other  genera),  on  Rosaceae  (Spiraea,  Rubus,  and  other 
genera),  and  on  a  few  other  plants.  A  few  of  the  local  Limnobiinae 
have  the  rostrum  very  greatly  elongated  — an  obvious  adaptation  for 
sucking  the  nectar  from  tubular  flowers.  The  species  of  Geranomyia 
feed  on  various  composite,  umbelliferous,  and  lauraceous  flowers. 
Toxorhina  frequents  composite,  rhamnaceous,  apocynaceous,  and  erica- 
ceous  plants.  The  exact  plants  frequented  are  discussed  under  the 
descriptions  of  the  respective  genera. 

Oviposition.—  The  females  lay  their  eggs  in  the  habitat  frequented  by 
the  larvae.  In  the  case  of  aquatic  forms  — as  Antocha,  Hexatoma, 
Eriocera,  and  other  genera  —  the  eggs  are  deposited  in  dipping  down  to 
the  water  surface,  one  or  more  eggs  being  deposited  at  each  descent. 
Forms  that  live  in  mud  or  moist  earth  lay  their  eggs  in  these  situations. 
Many  species  with  acute  ovipositors  insert  the  eggs  carefully  into  the  soil 
or  other  substance.  The  oviposition  of  Limnophila  (Eutonid)  alleni ^  as 
noted  by  the  writer  may  be  regarded  as  typical  of  this  class  of  species: 

Observations  made  at  Gloversville,  New  York,  June  28,  1916.  A  female  was  noted 
ovipositing  in  low,  wet  spots  along  a  small  woodland  stream.  She  flew  about  slowly  and 
silently,  just  skimming  the  ground,  until  a  place  suitable  for  egg-laying  was  found  She 
finally  chose  a  much-decayed  log  and  the  eggs  were  driven  home  securely  by  the  acicu^r 
tergal  valves  of  the  ovipositor.  Much  effort  is  expended  to  place  the  eggs  firmly  and  the 
rate  of  oviposition  is  not  more  than  eight  or  ten  a  minute,  the  female  often  pausing  to  rest 
for  ssveral  seconds.  While  thus  engaged,  the  fly  is  entirely  unconcerned  with  oti 
and  may  be  picked  up  by  hand. 

The  females  of  most  crane-flies  usually  live  but  a  short  time  after  egg- 
laying  is  completed.  Indeed,  the  entire  duration  of  life  of  the  adult 
crane-flies  is  probably  but  a  few  weeks  at  the  most. 


714 


CHARLES  PAUL  ALEXANDER 


The  specialized  methods  of  oviposition  in  the  Cylindrotominae  have 
already  been  discussed  on  page  709  under  the  account  of  Cylindrotoma 
splendens.  The  species  of  Tipula  and  otr^er  genera  the  females  of  which 
have  blunt  valves  to  the  ovipositor  (Styringomyia,  Macromastix,  and 
others) ,  all  probably  scatter  their  eggs  promiscuously  or  else  have  a  special- 
ized method  of  egg-laying.  Similarly,  the  species  of  Tipula  of  the  arctica 
group,  in  which  the  dorsal  valves  of  the  ovipositor  are  very  large,  placed 
horizontally,  and  with  the  margin  finely  serrated,  undoubtedly  have  a 
peculiar  method  of  oviposition,  but  this  has  not  yet  been  observed. 

The  egg 

The  number  of  eggs  laid  by  crane-flies  varies  from  about  forty-five  in 
Styringomyia  to  some  two  thousand  in  the  larger  species  of  Eriocera. 
The  following  table  indicates  the  general  range  in  the  group: 


Species 

Number  of  eggs  laid 

Reference 

Maximum 

Minimum 

Average 

Ptychoptera  albimana  
Styringomyia  didyma  
Eriocera  longicornis  
spinosa  
Hexatoma  megacera  
Phalacrocera  replicata  

587 
45 
1,034 
2,061 
372 
60 

60 
400 

297 
255 
602 
329 
366 
600 

520 

554 
45 
952 
1,942 
347 
60 

60 
300 
300 
289 
255 
602 
329 
309 
600 

Topsent,  1914-16 
Terry  ms. 
Alexander  ms. 
Alexander  ms. 
Alexander,  1915  c 
Miall  and  Shelford, 
1897 
Miiggenberg,  1901 
Lovett,  1915 
Hyslop,  1910 
Webster,  1893  b 
Webster,  1893  b 
Hyslop,  1910 
Alexander  ms. 
Alexander  ms. 
Del  Guercio,  1914 

872 
1,824 
316 

200 

'282 

251 

Liogma  glabrata  
Ctenophora  angustipennis 

Tipula  cunctans  

bicornis  
iephrocephala 

angustipennis  

collaris 

dejecta 

oleracea 

The  eggs  are  in  most  cases  elongate  with  the  ends  narrowed  and  rounded, 
or,  in  other  words,  spindle-shaped.  The  chorion  in  the  species  of  Tipula, 
Hexatoma,  and  other  genera  is  blackened  and  in  some  cases  more  or  less 
sculptured.  In  Ctenophora  angustipennis  (Lovett,  1915)  the  egg  is  ebony 
black  with  deep  purple  reflection,  elongate-oval,  uniform,  without  pits  or 
ridges,  and  measuring  from  1.26  to  1.4  millimeters  by  0.375  millimeter. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  715 

In  Tipula  bicornis  (Webster,  1893  b)  the  egg  is  0.8  millimeter  long  by  0.3 
to  0.4  millimeter  in  diameter,  elongate-ovoid,  with  one  side  deeply  concave, 
the  surface  highly  polished.  The  egg  of  Nephrotoma  ferruginea  is  smaller, 
with  five  distinct  grooves.  The  egg  of  Tipula  glacialis  (Pokorny,  1887:53) 
is  described  as  being  1.2  millimeters  long  and  only  0.3  millimeter  in  diame- 
ter, cylindrical,  the  two  ends  equally  rounded,  the  surface  smooth,  shiny 
black,  with  faint  steel-blue  or  purplish  red  reflections.  In  many  other 
genera  of  Tipulidae,  especially  the  smaller  forms,  the  eggs  are  soft  and 
whitish  or  nearly  hyaline.  In  a  few  species  the  eggs  take  on  a  decided 
green  or  greenish  tinge.  The  egg  of  Cylindrotoma  splendens  is  described 
elsewhere  in  this  paper  (page  709).  The  egg  of  Ptychoptera  albimana,  as 
described  by  Topsent  (1914-16),  measures  0.825  by  0.284  millimeter, 
and  is  pale  yellow,  slightly  arcuated,  and  with  the  surface  curiously 
ornamented. 

The  larva 

As  a  rule,  the  larvae  live  in  the  haunts  where  the  eggs  are  laid.  The 
duration  of  the  larval  stage  varies  from  about  a  month  in  Styringomyia 
(larva  and  pupa  together,  thirty-seven  days  —  Terry  ms.)  to  the  greater 
part  of  a  year  in  most  crane-flies.  Many  of  the  smaller  species  of  Eri- 
optera,  Ormosia,  Rhaphidolabis,  and  other  genera  are  on  the  wing  in  the 
spring  and  again  in  the  fall,  and  with  little  doubt  are  double-brooded. 
This  would  make  the  larval  existence  but  a  few  months,  but  still  probably 
longer  than  the  other  stages  taken  together.  This  problem  of  double 
broods  should  be  worked  out  carefully.  It  often  appears  that  there  are 
two  broods,  when  in  reality  there  may  be  two  developing  generations, 
each  passing  the  winter  as  larvae,  but  one  attaining  its  growth  much  more 
slowly  in  the  spring  and  summer  and  not  maturing  until  late  summer. 
Phalacrocera  and  Cylindrotoma  splendens  spend  about  eleven  months  in 
the  larval  stage  (Bengtsson  1897,  and  Cameron  1918),  while  Tipula 
paludosa  spends  nine  months  in  that  stage  (Rennie,  1917).  It  is  probable 
that  nearly  all  crane-flies  in  the  North  Temperate  Zone  winter  normally 
as  larvae.  The  growth  during  summer,  fall,  and  winter  is  very  slight,  but 
in  the  spring  it  is  greatly  accelerated  and  in  a  month  the  larva  may 
attain  its  full  growth.  Larvae  of  Tipula  ignobilis  taken  at  Ithaca,  New 
York,  on  April  23,  1917,  measured  only  6.5  millimeters  in  length;  on 
May  19  they  had  attained  their  full  growth  of  18  millimeters  and  were 
ready  to  pupate.  Cylindrotoma  splendens  spends  the  winter  as  a  larva, 


716  CHARLES  PAUL  ALEXANDER 

9  millimeters  in  length,  but  grows  rapidly  during  the  spring,  attaining 
its  full  size  (17  millimeters)  in  two  months  of  growth.  Liogma  glabraia 
spends  the  winter  as  a  very  small  larva,  but  in  the  spring  its  growth  is 
greatly  accelerated. 

The  haunts  in  which  the  larvae  of  crane-flies  occur  are  exceedingly 
varied.  In  the  case  of  single  large  genera,  such  as  Dicranomyia  and 
Tipula,  the  species  range  from  those  that  are  almost  strictly  aquatic  to 
others  that  are  entirely  terrestrial,  living  in  decaying  wood  or  even  mining 
in  the  leaves  of  plants. 

The  transition  between  strictly  aquatic  and  terrestrial  forms  is  very 
gradual,  as  was  pointed  out  by  Miall  (1895:11)  some  years  ago  when 
he  wrote: 

How  did  insects  ever  come  to  seek  the  water,  seeing  that  their  mode  of  respiration  is 
primarily  adapted  to  another  element?  We  can  see  almost  all  the  steps  of  the  adaptation 
on  the  shores  of  our  rivers,  lakes  and  seas.  We  can  see  dipterous  larvae  which,  like  the 
"  leather  jacket  "  (the  larva  of  the  daddy-long-legs),  burrow  in  the  ground  for  their  vegetable 
food,  and  devour  the  roots  of  grasses.  Other  larvae  of  the  same  family  (Tipulidae)  prefer 
moist  earth  in  the  neighborhood  of  streams.  Others  again  live  immersed  in  water,  or  mud 
saturated  with  water,  though  they  come  to  the  surface  at  times  and  push  their  tails,  which 
carry  the  spiracles,  into  the  air.  Some  few  have  become  so  completely  aquatic  that  they 
seldom,  if  ever,  come  to  the  surface,  and  all  their  supply  of  oxygen  is  obtained  from  the 
water. 

The  culmination  of  this  latter  condition  is  reached  in  forms  such  as 
Antocha  and  related  genera  and  species.  Crampton  (1919:100)  has 
made  similar  observations  on  the  subject. 

The  haunts  of  the  larvae  of  crane-flies  are  best  shown  by  the  following 
table,  in  which  the  various  species  are  arranged  according  to  habitat, 
from  the  strictly  aquatic  to  the  various  terrestrial  forms: 

Habitat  Species 

Strictly  aquatic,  in  silken  cases  Antocha 

In  very  rapid  water   (lotic)   on  or  in  sub-     Dicranomyia     simulans,     Pedicia,     Triogma, 
merged  mosses  (hygropetric  association)  Tipuline  No.  1,  and  others 

Aquatic,  on  submerged  plants  Phalacrocera,  Triogma 

Semi-aquatic   (part  of  life  spent  in  water,     Dicranomyia  simulans,  Eriocera,  Hexatoma, 
but  pupation  taking  place  on  land)  Aeshnasoma,     Longurio,     Tipula    abdomi- 

nalis,  T.  caloptera,  TV  bella,  and  others 

On  cliffs  and  wooden  walls,  usually  hi  silken     Dicranomyia  simulans,    Geranomyia,     Ellip- 
cases  covered  by  water  tera,  Dactylolabis,  and  others 

In  cold  springs  Pedicia,  Thaumastoptera 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II 


717 


Habitat    • 

In  stagnant  water  in  the  axils  of  bromelia- 
ceous  and  other  plants 

Amphibious  in  decaying  wood 

In  mud  or  sand: 

a.  In  open  swamps 


b.  In  shaded  woods 


c.  Along  the  margins  of  streams  and 
other  bodies  of  water 


In  or  beneath  damp  cushions  of  moss 

In  or  beneath  dry  cushions  of  moss 
In  dry  soil 

In  fungi 


In  decaying  vegetables,  plant  stems,  manure, 
and  like  situations 

In  wood: 

a.  In  decaying  wood,  usually  just  be- 
neath the  bark 


b.  In  nearly  solid  wood 

On  leaves  of  terrestrial  plants: 

a.  On  flowering  plants 

b.  On  mosses 

Mining  in  the  leaves  of  plants 


Species 
Trentepohlia,  Gnophomyia  rufa 

Protoplasa  (supp9sition) ,  Epiphragma 

Bittacomorpha,  Ptychoptera,  Rhamphidia, 
Erioptera  septemtrionis,  E.  chlorophylla,  E. 
vespertina,  Limnophila  adusta,  L.  macrocera, 
Pilaria  recondita,  P.  tenuipes,  P.  quadrata, 
Tricyphona  inconstans,  Prionocera,  Tipula 
dejecta,  T,  sayi,  T.  tricolor,  and  others 

Bittacomorphella,  Ormosia  innocens,  Eriop- 
tera megophthalma,  Molophilus  hirtipennis, 
Ulomorpha,  Dicranophragma,  Penthoptera, 
Tipula  cayuga,  and  others 

Limnobia  fallax,  Erioptera  (Hoplolabis) 
armata,  Trimicra,  Gonomyia  (Leiponeura) 
alexanderi,  G.  kansensis,  Eriocera,  Hexa- 
toma,  and  others 

Dicranomyia  badia,  D.  stulta,  Tipula  oro- 
pezoides,  T.  collaris,  T.  nobilis,  T.  ignobilis, 
and  others 

Liogma,  Dolichopeza,  Oropeza,  and  others 

Dicranoptycha,  Cladura,  Nephrotoma  ferrugi- 
nea,  Tipula  cunctans,  T.  angustipennis, 
and  others 

Limnobia  triocellata,  L.  cinctipes,  Ula,  and 
others 

Trichocera,  Limnobia  indigena,  Rhipidia 
domestica,  R.  maculata,  and  others 

Dicranomyia  rara,  D.  macateei,  Rhipidia 
fidelis,  R.  bryanti,  Elephantomyia,  Teucho- 
labis,  Gnophomyia,  Limnophila  unica, 
Brachypremna,  Ctenophora,  Dictenidia, 
Tipula  trivittata,  T.  usitata,  and  others 

Tanyptera 

Cylindrotoma 

Liogma,  Triogma  (in  some  instances) 

Dicranomyia  foliocuniculator 


Feeding. —  The  various  larvae  of  crane-flies  show  a  considerable  diversity 
in  their  habits  of  feeding.  The  majority  of  species,  as  known,  are  herbiv- 
orous, but  a  large  group  are  decidedly  carnivorous  in  their  habits. 

The  Tipulinae  feed  on  the  living  vegetable  tissue  or  plant  remains 
occurring  in  their  habitat,  or,  when  pressed  for  food,  they  will  eat  earth- 


718  CHAKLES  PAUL  ALEXANDER 

worms  (Patterson,  1908)  and  other  soft-bodied  animals.  Dissections  of 
Tipula  abdominalis  show  the  food  of  this  species  to  consist  principally  of 
small  filamentous  algae,  diatoms  (Diatoma,  Navicula,  and  others),  and 
rootlets  of  small  plants.  The  alimentary  tract  is  often  crammed  with 
sand  or  soil  particles.  The  species  of  Tipula  and  Nephrotoma  that  are 
injurious  to  plants,  considered  later  in  this  paper  under  the  heading 
Economic  Importance,  effect  their  damage  by  devouring  the  living  tissues 
of  the  roots.  The  Cylindrotominae  feed  on  the  living  tissues  of  the  plants 
on  which  they  dwell;  in  the  case  of  Phalacrocera,  Triogma,  and  Liogma 
these  are  mosses,  in  Cylindrotoma  they  are  the  parenchyma  and  epi- 
dermis of -higher  plants  (spermatophytes) .  With  this  habit  of  feeding 
on  plant  tissues  the  extreme  of  sluggishness  of  motion  is  attained.  The 
wood-boring  species  of  Tipulinae  (such  as  the  species  of  Tanyptera  and 
Ctenophora)  feed  on  the  ligneous  tissue  of  the  trees  hi  which  they  occur. 
Other  species  of  crane-flies  injure  young  seedlings  by  destroying  the  bark 
and  the  bast  tissues. 

The  hexatomine  and  pediciine  forms  represent  the  opposite  extreme, 
being  for  the  most  part  carnivorous  or  even  cannibalistic  in  their  habits. 
In  order  to  capture  their  prey  they  are  of  necessity  rapid  of  movement, 
and  in  this  group  the  most  graceful  and  active  of  all  tipulid  larvae  are 
found.  The  motions  of  the  species  are,  at  times,  exceedingly  agile  and 
snakelike.  The  food  consists  of  a  variety  of  animal  forms.  Dicranota 
has  been  recorded  as  feeding  on  worms  of  the  genus  Tubifex.  Pedicia 
usually  feeds  on  the  larvae  of  small  insects,  especially  Chironomidae, 
but  the  large  species  of  this  genus  and  of  Eriocera  are  capable  of  capturing 
almost  any  insect  of  a  size  equal  to  their  own.  The  larvae  of  Eriocera 
spinosa  are  able  to  inflict  painful  bites  on  tender  parts  of  the  skin  of  a  man. 
Penthoptera,  Eriocera,  Hexatoma,  and  most  of  the  limnophiline  groups 
likewise  feed  largely  on  midge  larvae.  From  one  small  larva  of  Limno- 
phila  (Dicranophragma)  fuscovaria  the  writer  has  dissected  out  the  remains 
of  two  large  midge  larvae,  whose  heads  were  nearly  half  the  size  of  that  of 
the  captor.  The  chitinized  mentum  in  these  predacious  forms  is  very 
weak  or  is  lacking,  allowing  for  a  tremendous  distension  of  the  gular  region. 
The  mandibles  are  always  developed  into  powerful  curved  hooks  which 
serve  well  their  purpose  of  grasping  and  holding  the  victims.  The 
esophageal  region  is  often  retrorsely  roughened  to  prevent  the  egress  of 
anything  that  has  once  started  down  the  throat.  In  addition  to  the 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  719 

various  chitinized  jaws,  legs,  heads,  and  other  insect  remains,  the  dis- 
tended proventricular  regions  usually  show  a  considerable  amount  of 
sand  particles  and  much  plant  tissue. 

The  pupa 

When  ready  to  pupate,  the  larva  ceases  feeding  and  becomes  much 
contracted  and  sluggish.  The  pupa  is  formed  within  the  last  larval  skin, 
which  is  then  shed  completely  except  in  certain  Cylindrotominae  and  a 
'ew  scattered  genera  in  other  tribes,  in  which  cases  the  larval  skin  adheres 
;o  the  posterior  end  of  the  abdomen.  The  pupal  existence  is  spent  in  or 
lear  the  haunts  of  the  larva. 

In  the  strictly  aquatic  genus  Antocha  the  pupa  lives  in  water  in  a  silken 
sase,  respiration  being  accomplished  by  means  of  the  many-branched 
wreathing  horns.  The  species  of  Elliptera  and  certain  Dicranomyia 
simulans,  for  example)  approach  this  aquatic  condition.  The  other 
pecies  of  crane-flies  with  aquatic  larvae  known  to  the  writer  go  to  the 
oil  in  order  to  pupate,  this  category  including  Eriocera,  Hexatoma,  Tip- 
da  caloptera,  T.  abdominalis,  and  many  others. 

The  pupae  of  some,  at  least,  of  the  Cylindrotominae  attach  themselves 

0  plant  stems  for  the  purpose  of  pupation.     The  leaf -mining  Dicrano- 
nyia  foliocuniculator  pupates  within  the  larval  passages.     The  majority 

1  the  limnobiine  forms  spend  the  pupal  existence  in  silken  cases  to  which 
3ebbles  and  particles  of  debris  or  plant  tissues  adhere. 

The  pupae  of  the  Ptychopteridae  have  one  of  the  two  breathing  horns 
enormously  elongated,  the  tip  of  this  being  projected  above  the  water 
evel  into  the  air  for  respiration.  Certain  tipuline  crane-flies  have  a  some- 
what similar  development  of  the  breathing  horns,  discussed  later. 

The  duration  of  the  pupal  existence  is  remarkably  uniform  thruout  the 
group,  averaging  from  six  to  eight  days.  The  following  table  illustrates 
this  for  the  more  representative  genera  and  species.  Records  which 
have  not  been  determined  sufficiently  close,  and  which  as  stated  are 
probably  too  long,  are  indicated  by  an  asterisk. 


720 


CHARLES  PAUL  ALEXANDER 


Species 

Duration  of  pupal  .existence  (days) 

Reference 

Maximum 

Minimum 

Average 

Ptychoptera  rufocincta. 

10  * 

Alexander  ms. 
Topsent,  1914-16 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Shannon  ms. 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Alexander,  19  15  a 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Beling,  1879:46 
Alexander  ms. 
Alexander  ms. 
Hyslop  ms. 
Alexander  ms. 
Johnson  ms. 
Shannon  ms. 
Cameron,  1918 
Miiggenberg,  1901 
Bengtsson,  1897 
Beling,  1886  (assylvicok 
Lovett,  1915 
Beling,  1873  b 
Beling,  1886 
DelGuercio,  1914:315 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Alexander  ms. 
Beling,  1879  (as  histru 
Beling,  1879 
Beling,  1886 

albimana.. 

*12 
*13 

7 
*8 

7 

.11 

Bittacomorphella  . 

Dicranomyia  badia 

:  oo  01- 

7 
..„. 

stulta 

Rhipidia  bryanti  . 

Limnobia  cinctipes 

triocellata 

9 
6 
*10 
*10 

84 
6 

Rhamphidia  mainensis 

Dicranoptycha  winnemana  
Ula  elegans 

Epiphragma 

6 

8 
8 
8 

7 

HNH|N 

CO  GO  GO  O  1>- 

Pseudolimnophila 

9 
9 
*14 

7 
*9 
7 
7.3 

*142 
-8 
*10 
5 

9 
8 
10 
11 
8 
6 

•y 

*14 

8 
7.6 
8 
8 
8 
7 
7 
*12 

Limnophila  macrocera 

fuscovaria  

Pilaria  

Ulomorpha  . 

Eriocera  fultonensis  
longicornis 

7 

7 
7.3 
64 

104 

"7 

8 

74 
6 
10 
7 
*12 

74 

8 

74 

7 
*10 

Hexatoma  megacera  

"7 
7 

4 

6 

6 
10 

7 

"*io" 

7 
"7' 
"7' 

"*8" 

Pedicia  rivosa  

Erioptera  macrophthalma 

Molophilus  hirtipennis  

Gnophomyia  tristissima 

Ormosia  nigripila  

Teucholabis  complexa  

Elephantomyia  
Cylindrotoma  splendens  
Liogma  glabrata  

Phalacrocera  replicata  
Dolichopeza  albipes 

Ctenophora  angustipennis  
Dictenidia  bimaculata 

Tipula  maxima  . 

oleracea  . 

cayuaa  .  , 

ignobilis    . 

dejecta  
collaris  
Nephrotoma  lineata  
lunulicornis  
analis  .  . 

THE  CKANE-FLIES  OF  NEW  YORK  —  PART  II  721 

ENEMIES 

Crane-flies  have  many  enemies  and  but  few  means  of  combating  them. 
Every  stage  of  the  crane-fly's  existence  is  fraught  with  danger.  The 
larvae  of  the  majority  of  species  are  soft-bodied,  herbivorous  creatures, 
which  form  a  choice  morsel  of  food  for  carnivorous  forms  of  many  classes 
and  orders.  The  period  when  the  adult  fly  has  just  emerged  from  the 
pupal  hull  is  undoubtedly  the  one  in  which  the  greatest  danger  is  found, 
for  then  the  insects  are  teneral  and  incapable  of  rapid  motion.  The 
adult  flies  are  sometimes  drowned  in  the  sea  or  other  large  bodies  of  water 
and  their  dead  fragments  cast  up  in  windrows  on  the  shore.  A  very 
unusual  instance  of  this  kind  is  recorded  for  Tipula  oleracea  by  Patterson 
(1908).  At  their  best,  the  adult  flies  are  poor,  awkward  fliers  and  are 
easily  captured  by  a  wide  range  of  species,  as  indicated  in  the  following 
pages. 

Predatory  natural  enemies 
Vertebrates 

Mammalia.—  Undoubtedly  many  of  the  smaller  mammals  prey  on  the 
larvae  of  various  species  of  Tipula.  In  another  paper  (Alexander,  1919  d: 
776-777)  the  writer  has  discussed  the  value  of  the  larvae  of  an  undeter- 
mined species  of  Tipula  in  the  Pribilof  Islands  as  an  article  of  food  for  the 
arctic  fox.  These  larvae  occur  in  enormous  numbers  beneath  the  lichens 
of  the  tundra  and  the  foxes  can  easily  get  them  at  a  time  when  other  food 
is  scarce  or  unobtainable.  Mice,  shrews,  and  moles  find  an  important 
element  of  their  food  from  this  source.  White  (1914)  states  that  in  North 
Wales  the  European  mole,  Talpa  europaea,  eats,  on  an  average,  twenty 
crane-fly  larvae  a  day,  these  constituting  one  of  the  main  foods  of  this 
mammal. 

Aves.—  Birds  are  well-known  enemies  of  crane-flies,  both  in  their  imma- 
ture stages  and  as  adult  flies.  Hyslop  (1910:129-130)  lists  ninety-one 
species  of  birds  which  are  known  to  feed  on  crane-flies.  Baer  (1913) 
describes  ravens  and  starlings  as  feeding  on  the  larvae  of  a  species  of 
Tipula.  While  working  with  the  United  States  Biological  Survey,  the 
writer  was  given  the  opportunity  to  examine  files  for  records.  His  thanks 
for  assistance  in  this  work  are  due  to  Messrs.  Kalmbach,  McAtee,  and 
Wetmore.  The  records  of  the  Survey  are  based  on  an  examination  of 
the  contents  of  the  stomachs  of  many  thousands  of  individuals,  and 
furnish  very  valuable  and  fairly  complete  data  on  the  North  American 
2 


722  CHARLES  PAUL  ALEXANDER 

birds  which  prey  .on  various  stages  of  the  Tipulidae.  These  records  are 
here  arranged  according  to  the  latest  check  list  of  the  American  Orni- 
thologists' Union : 8 

Longipennes: 
Laridae: 

Larus  delawarensis  Ord.     Ring-billed  gull 
L.  franklinii  Rich.     Franklin's  gull 
Sterna  hirundo  Linn.     Common  tern 
Anseres:  / 

Anatidae: 

Mergus  americanus  Cass.     Merganser 
Anas  platyrhynchos  Linn.     Mallard 
A.  rubripes  Brewst.     Black  duck 
Mareca  americana  (Gmel.).     Baldpate 
Nettion  carolinense  (Gmel.).     Green-winged  teal 
Dafila  acuta  (Linn.).     Pintail 
Aix  sponsa  (Linn.).    Wood  duck 
Herodiones: 
Ardeidae: 

Egretta  candidissima  candidissima  (Gmel.) .     Snowy  egret 
Butorides  virescens  virescens  (Linn.).     Green  heron 
Limicolae: 

Phalaropodidae : 

Phalaropus  fulicarius  (Linn.).     Red  phalarope 
Lobipes  lobatus  (Linn.).     Northern  phalarope 
Steganopus  tricolor  Vieill.     Wilson's  phalarope 
Recurvirostridae : 

Recurvirostra  americana  Gmel.     Avocet 
Himantopus  mexicanus  (Mull.).     Black-necked  stilt 
Scolopacidae: 

Philohela  minor  (Gmel.).     American  woodcock 
Gallinago  delicata  (Ord).     Wilson's  snipe 
Arquatella  maritima  maritima  (Briinn.).     Purple  sandpiper 
Pisobia  aurita  (Lath.).     Sharp-tailed  sandpiper 
P.  maculata  (Vieill.).     Pectoral  sandpiper 
P.  bairdii  (Coues).     Baird's  sandpiper 
Limosafedoa  (Linn.).     Marbled  godwit 
Totanus  flavipes  (Gmel.).     Yellow-legs 
Heteractitis  incanus  (Gmel.).     Wandering  tatler 
Bartramia  longicauda  (Bechst.).     Bartramian  sandpiper 
Charadriidae: 

Charadrius  dominicus  dominicus  (Miill.).     Golden  plover 
Oxyechus  vociferus  (Linn.).     Killdeer 
Aphrizidae: 

Arenaria  inter pres  inter pres  (Linn.).     Turnstone 
Gallinae: 

Tetraonidae: 

Bonasa  umbeUus  umbellus  (Linn.).     Ruffed  grouse 
Raptores: 

Buteonidae: 

Ictinia  mississippiensis  (Wils.).     Mississippi  kite 


3  Check  list  of  North  American  birds,  3d  ed.,  p.  1-430.     American  Ornithologists'  Union.     1910, 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  723 

Hoccyges: 

Cuculidae: 

Coccyzus  americanus  americanus  (Linn.).     Yellow-billed  cuckoo 

C.  erythrophthalmus  (Wils.).     Black-billed  cuckoo 
Cuculus  canorus  telephonus  Heine.     Kamchatka  cuckoo 

'ici: 

Picidae: 

Dry  abates  pubescens  pubescens  (Linn.).     Downy  woodpecker 

D.  nuttallii  (Gamb.).     Nuttall's  woodpecker 
Sphyrapicus  varius  varius  (Linn.).     Yellow-bellied  sapsucker 
S.  thyroideus  (Cass.).     Williamson's  sapsucker 
Melanerpes  erythrocephalus  (Linn.).     Red-headed  woodpecker 
Asyndesmus  lewisi  Riley.     Lewis's  woodpecker 

Colaptes  auratus  auratus  (Linn.).     Flicker 
Vlacrochires: 

Capri  mulgidae: 

Antrostomus  carolinensis  (Gmel.).     Chuck-will's-widow 

Chordeiles  virginianus  virginianus  (Gmel.).     Nighthawk 

C.  acutipennis  texensis  Lawr.     Texas  nighthawk 
Cypselidae: 

Chaetura  pelagica  (Linn.J.     Chimney  swift 

C.  vauxi  (J.  K.  Towns.).     Vaux's  swift 

Aeronautes  melanoleucus  (Baird).     White-throated  swift 
Trochilidae: 

Calypte  anna  (Less.).     Anna's  humming  bird 

Selasphorus  rufus  (Gmel.).     Rufous  humming  bird 
asseres: 

Tyrannidae: 

Tyrannus  tyrannus  (Linn.).     Kingbird 

T.  verticalis  Say.     Arkansas  kingbird 

T.  vociferans  Swains.     Cassin's  kingbird 

Myiarchus  crinitus  (Linn.).     Great-crested  flycatcher 

Sayornis  phoebe  (Lath.).     Phoebe 

S.  sayus  (Bonap.).     Say's  phoebe 

S.  nigricans  (Swains.).     Black  phoebe 

Myiochanes  virens  (Linn.).     Wood  pewee 

M.  richardsonii  richardsonii  (Swains.).     Western  wood  pewee 

Empidonax  flaviventris  (W.  M.  &  S.  F.  Baird).     Yellow-bellied  flycatcher 

E.  difficihs  difficilis  Baird.     Western  flycatcher 

E.  virescens  (Vieill.).     Acadian  flycatcher 

E.  traillii  traillii  (Aud.).     Traill's  flycatcher 

E.  minimus  (W.  M.  &  S.  F.  Baird).     Least  flycatcher 

E.  wrightii  Baird.     Wright's  flycatcher 

E.  griseus  Brewst.     Gray  flycatcher 
Corvidae: 

Pica  pica  hudsonia  (Sab.).     American  magpie 

Cyanocitta  cristata  cristata  (Linn.).     Blue  jay 

C.  stelleri  stelleri  (Gmel.).     Steller's  jay    ' 

Corvus  brachyrhynchos  brachyrhynchos  Brehm.     American  crow 

C.  ossifragus  Wils.     Fish  crow 
Sturnidae: 

Sturnus  vulgaris  Linn.     Starling 
Icteridae: 

Dolichonyx  oryzivorus  (Linn.).     Bobolink 

Molothrus  ater  ater  (Bodd.).     Cowbird 


724  CHARLES  PAUL  ALEXANDER 

Passeres  (continued): 

Icteridae  (continued) : 

Xanthocephalus  xanthocephalus  (Bonap.).     Yellow-headed  blackbird 

Agelaius  phoeniceus  phoeniceus  (Linn.)-     Red-winged  blackbird 

A.  gubernator  californicus  Nels.     Bicolored  redwing 

Sturnella  magna  magna  (Linn.).     Meadow  lark 

Icterus  spurius  (Linn.).     Orchard  oriole 

/.  galbula  (Linn.).     Baltimore  oriole 

/.  bullockii  (Swains.).     Bullock's  oriole 

Euphagus  cyanocephalus  (Wagl.).     Brewer's  blackbird 

Quiscalus  quiscula  quiscula  (Linn.).     Purple  grackle 
Fringillidae: 

Leucosticte  griseoriucha  (Brandt).     Aleutian  rosy  finch 

Plectrophenax  nivalis  nivalis  (Linn.).     Snow  bunting 

P.  hyperboreus  Ridgw.     McKay's  snow  bunting 

Calcarius  lapponicus  lapponicus  (Linn.).     Lapland  longspur 

Passer culus  sandwichensis  savanna  (Wils.).     Savanna  sparrow 

Zonotrichia  leucophrys  leucophrys  (J.  R.  Forst.).     White-crowned  sparrow 

Z.  albicollis  (Gmel.).     White-throated  sparrow 

Junco  hyemalis  hy emails  (Linn.).     Slate-colored  junco 

Passer  domesticus  (Linn.).     English  sparrow 

Melospiza  melodia  melodia  (Wils.).     Song  sparrow 

M.  lincolnii  lincolnii  (Aud.).     Lincohi's  sparrow 

M.  georgiana  (Lath.).     Swamp  sparrow 

Passerella  iliaca  iliaca  (Merr.).     Fox  sparrow 

Pipilo  erythrophthalmus  erythrophthalmus  (Linn.).     Towhee 

Passerina  amoena  (Say).     Lazuli  bunting 
Tangaridae: 

Piranga  ludoviciana  (Wils.).     Western  tanager 

P.  erythromelas  (Vieill.).     Scarlet  tanager 

P.  rubra  rubra  (Linn.).     Summer  tanager 
Hirundinidae: 

Progne  subis  subis  (Linn.).     Purple  martin 

Petrochelidon  lunifrons  lunifrons  (Say).     Cliff  swallow 

Hirundo  erythrogastra  Bodd.     Barn  swallow 

Iridoprocne  bicolor  (Vieill.).     Tree  swallow 

Tachycineta  thalassina  lepida  Mearns.     Northern  violet-green  swallow 

Riparia  riparia  (Linn.).     Bank  swallow 

Stelgidopteryx  serripennis  (Aud.).     Rough-winged  swallow 
Bombycillidae: 

Bombycilla  cedrorum  Vieill.     Cedar  waxwing 
Vireonidae: 

Vireosylva  olivacea  (Linn.).     Red-eyed  vireo 

V.  philadelphica  (Cass.).     Philadelphia  vireo 

V.  gilva  gilva  (Vieill.).     Warbling  vireo 

Lanivireo  solitarius  solitarius  (Wils.).     Blue-headed  vireo 

Vireo  griseus  griseus  (Bodd.).     White-eyed  vireo 

V.  bellii  bellii  Aud.     Bell's  vireo 

V.  bellii  pusillus  Coues.     Least  vireo 
Mniotiltidae: 

Mniotilta  varia  (Linn.).     Black-and-white  warbler 

Helmitheros  vermivorus  (Gmel.).     Worm-eating  warbler 

Dendroica  aestiva  aestiva  (Gmel.).     Yellow  warbler 

D.  coronata  (Linn.).     Myrtle  warbler 

D.  auduboni  auduboni  (Towns.).    Audubon's  warbler 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  725 

Passeres  (continued) : 

Mniotiltidae  (continued) : 

D.  dominica  dominica  (Linn.).     Yellow-throated  warbler 

Oporornis  tolmiei  (J.  K.  Towns.).     Macgillivray's  warbler 

Geothlypis  trichas  trichas  (Linn.).     Maryland  yellowthroat 

Icteria  virens  virens  (Linn.).     Yellow-breasted  chat 

Wilsonia  pusilla  pusilla  (Wils.).     Wilson's  warbler 

Setophaga  ruticilla  (Linn.).     American  redstart 
Motacillidae: 

Anthus  rubescens  (Tunst.).     Pipit 
Mimidae: 

Mimus  polyglottos  polyglottos  (Linn.).     Mocking  bird 

Dumetella  carolinensis  (Linn.).     Catbird 

Toxostoma  rufum  (Linn.).     Brown  thrasher 

T.  redivivum  (Gamb.).     Calif ornian  thrasher 
Troglodytidae: 

Heleodytes  brunneicapillus  couesi  (Sharpe).     Cactus  wren 

Salpinctes  obsoletus  obsoletus  (Say).     Rock  wren 

Catherpes  mexicanus  punctulatus  Ridgw.     Dotted  canon  wren 

Thryothorus  ludovicianus  ludovicianus  (Lath.).     Carolina  wren 

Thryomanes  bewickii  bewickii  (Aud.).     Bewick's  wren 

Nannus  hiemalis  hiemalis  (Vieill.).     Winter  wren 

Telmatodytes  palustris  palustris  (Wils.).     Long-billed  marsh  wren 
Paridae: 

Baeolophus  inornatus  inornatus  (Gamb.).     Plain  titmouse 

Penthestes  atricapillus  atricapillus  (Linn.).     Chickadee 

P.  carolinensis  carolinensis,  (Aud.) .     Carolina  chickadee 

P.  gambeli  gambeli  (Ridgw.).     Mountain  chickadee 

P.  hudsonicus  hudsonicus  (Forst.).     Hudsonian  chickadee 

Psaltriparus  minimus  minimus  (J.  K.  Towns.).     Bush-tit 
Chamaeidae: 

Chamaea  fasciata  fasciata  (Gamb.).     Wren-tit 
Sylviidae: 

Regulus  calendula  calendula  (Linn.).     Ruby-crowned  kinglet 
Turdidae: 

Myadestes  townsendi  (Aud.).     Townsend's  solitaire 

Hylocichla  mustelina  (Gmel.).     Wood  thrush 

H.  fuscescens  fuscescens  (Steph.).     Veery 

H.  aliciae  aliciae  (Baird).     Gray-cheeked  thrush 

H.  ustulata  ustulata  (Nutt.).     Russet-backed  thrush 

H.  ustulata  swainsoni  (Tschudi).     Olive-backed  thrush 

H.  guttaia  pallasii  (Cab.).     Hermit  thrush 

Planesticus  migratorius  migratorius  (Linn.).     American  robin 

The  principal  families  that  feed  on  the  adult  flies  are  the  Caprimulgidae, 
the  Cypselidae,  the  Tyrannidae,  the  Icteridae,  the  Hirundinidae,  the 
Mniotiltidae,  the  Troglodytidae,  the  Paridae,  and  the  Turdidae. 

Many  species  of  birds  feed  on  the  larvae  of  crane-flies,  the  more  notable 
of  these  being  the  water-fowl,  Anatidae,  the  shore  birds,  Scolopacidae, 
and  the  thrushes,  Turdidae.  Sim  (1907)  has  recorded  the  chestnut- 
sided  warbler,  Dendroica  pensylvanica  (Linn.)  as  feeding  on  Tipula  sp. 


726  CHARLES  PAUL  ALEXANDER 

In  Europe  the  raven  and  the  starling  are  important,  as  already  stated. 
Patterson  (1908)  records  the  starling  as  eating  great  numbers  of  Tipula 
oleracea  and  as  boring  into  the  soil  in  search  of  the  larvae  of  this 
species. 

It  will  be  noted  that  in  the  foregoing  list  no  species  of  doves  or  pigeons 
are  recorded  in  this  country  as  feeding  on  Tipulidae.  The  following  note 
on  an  Australian  pigeon  shows  the  importance  of  the  larvae  as  a  food  for 
these  birds: 4 

Mr.  North  exhibited  the  head,  crop,  and  gizzard  of  a  wonga-wonga  pigeon  (Leucosarcia 
picata  Lath.)  shot  by  Mr.  H.  J.  McCooey  in  a  myrtle  scrub  at  Upper  Burragorang  on  the 
21st  inst.  The  crop  is  absolutely  crammed  with  dipterous  larvae  (Habromastix  cinerascens 
Sk.)  and  undigested  portions  of  them  mixed  with  seeds,  berries,  and  earth  appear  also  in  the 
,  gizzard.  As  the  larvae  are  known  to  be  destructive  to  grass,  the  wonga-wonga  would  appear 
to  be  deserving  of  consideration. 

Amphibia. —  Crane-flies,  both  larvae  and  adults,  form  a  considerable 
element  of  the  food  of  many  Amphibia.  The  studies  by  Needham 
(1905 : 13)  show  this  to  be  true  in  the  case  of  the  bullfrog,  Rana  cates- 
beiana  Shaw.  Munz  (1920)  studied  the  food  habits  of  eight  species  of 
Anura  and  found  that  five  fed  on  crane-fly  larvae  or  adults.  These 
species  were  Rana  damitans  Latr.,  R,  sylvatica  Lee.,  R.  palustris  Lee., 
R.  pipiens  Schreb.,  and  Hyla  crucifer  Wied.  The  following  additional 
records  are  given: 

Desmognathus  fusca  Raf.  Dusky  salamander.  Wings  of  a  Limnophila  found  in  a  speci- 
men from  Ithaca,  New  York.  (A.  A.  Noyes.) 

Rana  damitans  Latr.  Green  frog.  Two  larvae  of  a  Tipula,  near  dejecta  Walker,  found 
in  stomach.  (S.  W.  Frost.) 

Bufo  lentiginosus  woodhousei  Girard.  Centerville,  Utah,  April  27,  1912.  Twenty-six 
per  cent  of  the  food  of  this  species  consisted  of  a  large  tipulid.  (E.  R.  Kalmbach.) 

Bufo  sp.  Washington,  D.  C.,  May  17,  1890.  One  per  cent  tipulid  larvae.  (W.  L. 
McAtee.) 

Pisces. —  The  larvae  of  crane-flies  furnish  favorite  morsels  for  many 
carnivorous  species  of  fish,  and  as  a  consequence  they  are  in  considerable 
demand  with  fishermen  as  bait  for  bass  and  other  game  fish.  These 
include  the  larvae  of  the  larger  species  of  Eriocera  and  many  large  semi- 
aquatic  species  of  Tipula,  especially  Tipula  caloptera  and  T.  abdominalis. 
Fragments  of  the  adult  flies  are  often  found  in  the  stomach  contents  of 
fish,  notably  species  of  trout,  most  of  these  pieces  being  of  individuals  that 


4  Linn.  Soc.  New  South  Wales.     Proc.,  ser.  2:g:585.     1894. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  727 

. 

had  fallen  into  the  water  or  were  captured  while  newly  transformed. 
Forbes  (1888)  records  Coregonus  and  Hyodon  as  feeding  on  the  eggs, 
and  Notropis  as  feeding  on  the  larvae,  of  crane-flies.  Levander  (1909:1) 
records  Perca  fluviatilis  Linn,  as  feeding  on  larvae  of  Tipulidae. 

Invertebrates 

Arachnida. —  Some  spiders  are  notable  enemies  of  crane-flies,  which 
fall  easy  victims  while  in  a  helpless,  teneral  condition.  These  spiders 
represent  many  families,  such  as  the  Thomisidae,  the  Lycosidae,  the 
Attidae,  the  Epeiridae,  and  others  (Alexander  and  Lloyd,  1914:15; 
also,  Alexander,  1915  c:  144).  An  account  of  their  preying  on.  the  larger 
crane-flies  of  the  genus  Tipula  is  given  under  the  discussion  of  T. 
taughannock  (page  1013).  In  a  recent  paper  Bilsing  (1920)  has  recorded 
four  species  of  crane-flies  eaten  by  spiders.  These  spiders  represented 
thirteen  species  arranged  in  four  families  —  the  Lycosidae,  the  Attidae, 
the  Epeiridae,  and  the  Agelenidae.  J.  R.  Malloch  found  a  female  Tipula 
angustipennis  which  was  being  eaten. by  a  crab  spider,  determined  by 
Mr.  Banks  as  Xysticus  ferox  Htz.  .The  spider  did  not  release  its  hold 
until  after  it  was  placed  in  a  jar  of  cyanide. 

Hexapoda. —  Odonata:  The  following  records  of  dragon-flies  found 
feeding  on  adult  crane-flies  are  available:  Gomphus  vastus  Walsh  and 
G.  ventricosus  Walsh,  found  feeding  on  Tipula  bicornis  at  South  Bend, 
Indiana,  May  30,  1914  (E.  B.  Williamson);  Helocordulia  uhleri  (Selys), 
found  feeding  on  Eriocera  longicornis  (Alexander,  1915 c:  152);  Agrion 
puella  Linn.,  found  feeding  on  Erioptera  flavescens  (Campion,  1914:498). 
C.  H.  Kennedy  has  sent  the  writer  a  male  and  a  female  specimen  of  a  small 
crane-fly,  Teucholabis  pabulatoria  Alex.,  which  he  found  in  the  mouth  of 
a  damsel  fly,  Hetaerina  tricolor  Burm.,  collected  in  Guatemala  by  Professor 
Hine.  Needham  and  Hart  (1901-03  [1901]:  47)  record  the  nymphs  of 
Anax  junius  (Dm.)  feeding  on  the  larvae  of  Tipulidae. 

Diptera:  The  adult  flies  of  at  least  three  families  of  the  order  Diptera 
— the  Asilidae,  the  Empididae,  and  the  Scatophagidae — and  the  larvae 
of  a  fourth,  Anthomyiidae,  are  notable  enemies  of  crane-flies.  The 
records  of  Kirby  (1892),  Poulton  (1906-07),  Bromley  (1914),  Alexander 
(1915 c),  and  McAtee  and  Banks  (1920)  on  the  first  three  of  these 
families  may  be  summarized  as  follows: 


728  CHARLES  PAUL  ALEXANDER 

Asilidae  (robber  flies)  Prey 

Lasiopogon  cinctus  (Fabr.)  Nephrotoma  lineata  (Scop.)  (Poulton,  1906-07, 

as  histrio  [Fabr.]) 
Neoitamuscyanurus  (Lw.)  (in  copulation  with     Tipula  scripta  Meig.  (Poulton,   1906-07) 

female,  the  latter  with  the  crane-fly) 

Proctacanthus  philadelphicus  Macq.  Nephrotoma  sp.  (2  records,  Bromley,  1914) 

Asilus  sp.  Nephrotoma   ferruginea   (Fabr.)    (3  records, 

Alexander,  1915  c) 

Asilus  flavofemoratus  Hine  Tipula  sp.  (McAtee  and  Banks,   1920:30) 

Asilus  notatus  Wied.  Nephrotoma  ferruginea  (Fabr.)  (McAtee  and 

Banks,  1920:31) 
Asilus  paropus  Walk.  Epiphragma   solatrix    (O.    S.)    (McAtee   and 

Banks,  1920:31) 
Asilinae  Tipula  lateralis  Meig.  (Poulton,  1906-07) 

The  Empididae,  or  dance  flies,  are  small,  predacious  flies  which  are  closely  related  to  the 
robber  flies  and  largely  replace  them  in  moist  shaded  situations.  They  unquestionably 
play  a  most  important  part  in  the  economy  of  the  Tipulidae,  since  both  groups  are  practically 
confined  to  the  same  general  situations  and,  in  the  temperate  regions  at  least,  representatives 
of  the  two  groups  are  invariably  found  together.  Macquart  (Kirby,  1892:229),  discussing 
Empis  livida  Linn.,  writes  as  follows:  "  Among  the  thousands  of  pairs  which  I  have  noticed 
resting  on  bushes  or  hedges,  nearly  all  the  females  were  engaged  in  sucking  some  insect, 
sometimes  small  Phryganidae  or  Ephemeridae,  but  more  often  Tipulidae.  They  busy 
themselves  with  feeding  and  perpetuating  their  species  at  the  same  time."  Hewlett  (1907) 
records  Empis  borealis  Linn,  as  feeding  on  Tipulidae.  The  following  records  are  from 
Poulton  (1906-07:380-382):  Empis  tesselata  Fabr.  preys  on  Tipula  lunata  Linn.,  T. 
paludosa  Meig.,  and  other  species;  Empis  livida  Linn,  preys  on  Dicranomyia  sp.  (It  is 
probable  that  the  long-legged  flies,  Dolichopodidae,  likewise  play  an  important  r61e  in 
the  lives  of  the  Tipulidae,  but  no  records  are  available  to  the  writer  to  confirm  this  belief.) 

The  Scatophagidae  (Cordyluridae) ,  or  dung  flies,  unquestionably  play  an  important  part 
hi  the  lives  of  the  smaller  crane-flies  (Limnobiinae).  The  following  records  indicate  this 
relationship,  the  last  being  supplied  by  Malloch: 

Enemy  Prey 

Scatophaga  suilla  (Fabr.)  Dicranomyia  lutea  (Meig.)  (Poulton,  1906-07) 

Scatophaga  sp.  (stercoraria?)  Erioptera  sp.  (Poulton,  1906-07) 

Scatophaga  sp.  Hexatoma  megacera  (O.  S.)  (Alexander,  1915  c^ 

Scatophag aster cor aria (L.)  and  squalida Meig.  Trichocera  (Malloch,  1911) 

Comparatively  recently  the  larvae  of  Anthomyiidae  have  been  found 
to  play  a  highly  important  role  in  the  economy  of  other  insects  living  in 
the  same  haunts.  The  very  important  paper  by  Keilin  (1917)  may  be 
consulted  in  connection  with  this  point.  This  student  found  the  follow- 
ing Anthomyiidae  associated  with  Tipulidae:  Graphomyia  maculata  Scop., 
feeding  on  larvae  of  Ptychoptera  contaminata  (L.)  (Keilin,  1917:354-360); 
Allognota  agromyzina  Fall.,  associated  with  and  possibly  feeding  on  the 
larvae  of  Ula  macroptera  Macq.  (page  380-362  of  reference  cited); 
Phaonia  cincta  Zett.,  feeding  on  larvae  and  pupae  of  Mycetobia  pallipes 
(page  362-375  of  reference);  Phaonia  goberti  Mik,  associated  with  and 
possibly  feeding  on  the  larvae  of  Gnophomyia  tripudians  Bergr.  (page 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  729 

375-377  of  reference);  Mydaea  pertusa  Meig.,  possibly  feeding  on  larvae 
of  Tipulidae  (page  393-396  of  reference). 

A  species  of  Phaonia  with  presumably  parasitic  habits  is  discussed  on 
page  732  of  this  memoir. 

Coleoptera :  The  adults  and  the  larvae  of  the  following  species  of  ground 
beetles  (Carabidae)  have  been  recorded  as  important  enemies  of  crane- 
flies  (Hyslop,  1910) :  Poedlus  lucublandus  Say;  Micromaseus  femoralis 
(Kirby);  Platynus  sp.;  Harpalus  pennsylvanicus  Dej.;  H.  caliginosus  Fabr. 
Most  of  these  were  recorded  by  Webster  (1893  a: 241)  as  feeding  on  the 
injurious  crane-fly  Tipula  bicornis. 

The  larvae  of  Elateridae  (wireworms)  are  enemies  of  crane-flies.  The 
writer  has  seen  several  feeding  on  large  larvae  of  Tipula  trivittata. 

Hymenoptera:  Ants  (Formicidae)  undoubtedly  capture  and  devour 
many  crane-flies,  these  generally  being  pupae  or  teneral  adults.  Hyslop 
(1910)  cites  the  record  of  an  Aphaenogaster  fulva  Roger  which  was  observed 
dragging  a  living  adult  tipulid  over  the  ground.  Chapman  (1918:191) 
records  feeding  species  of  Myrmica  on  adult  Tipulidae  which  were  readily 
eaten  by  the  ants  in  preference  to  most  other  insect  food.  On  August  2, 
1917,  at  Lamed,  Pawnee  County,  Kansas,  along  the  bank  of  the  Arkansas 
River,  the  writer  noted  a  small  ant  which  was  dragging  a  still  living 
pupa  of  Gonomyia  kansensis  from  its  burrow  in  the  sand  at  the  bank  of 
the  river.  The  ant  had  the  pupa  about  the  head  and  carried  it  off  despite 
its  struggles.  Both  the  Gonomyia  and  the  formicid  were  common  and 
the  tragedy  as  described  was  not  'accidental. 

The  Pemphredonidae  (Mimesinae)  and  the  Crabronidae,  fossorial  wasps 
in  the  Hawaiian  Islands,  bear  an  important  relationship  to  the  smaller 
Tipulidae  dwelling  in  the  same  haunts.  Writing  of  the  Mimesinae,  a 
subfamily  of  the  Pemphredonidae,  Perkins  (1913:lxxxv)  says: 

All  the  Hawaiian  species  of  both  genera  are  true  forest  insects  and  most  of  them  may  be 
seen  in  large  numbers,  where  they  occur,  flying  around  ferns  and  bushes  in  sunny  places. 
The  males  are  often  much  more  numerous  apparently  than  the  females,  but  this  is  due  to 
the  more  retiring  habits  of  the  latter,  which,  when  they  have  begun  to  provision  their  nests, 
frequent  dark,  shady  and  damp  places  in  search  of  their  prey.  This  consists  of  the  endemic 
Limnobiidae  or  daddy  longlegs,  which  live  in  such  places.  .  .  .  Although  I  have  often 
watched  females  of  these  wasps  returning  with  prey  to  their  burrows,  it  was  always  Tipulidae 
that  they  carried  to  the  nest.  Their  burrows  are  usually  made  in  the  ground  and  are  often 
drilled  down  from  beneath  a  stone,  this  no  doubt  serving  to  keep  the  burrow  sufficiently 
dry. 


730  CHAELES  PAUL  ALEXANDER 

Referring  to  Limnobiinae,  Perkins  says  (page  clxxxii  of  the  same 
reference) : 

The  species  of  Dicranomyia  are  endemic,  and  to  the  five  described,  others  and  perhaps 
many  more  will,  no  doubt,  be  added.  They  are  common  insects,  sometimes  attracted  by 
light  and  sometimes  observed  at  rest  in  the  daytime,  or  on  the  wing  in  the  dark  cavity  of 
some  hollow  tree.  In  the  latter  case  a  number  are  usually  seen  flying  together,  rising  and 
falling  in  their  flight  in  the  narrow  space  of  a  few  feet.  We  have  bred  one  or  more  species 
from  decayed  wood,  overgrown  with  damp  moss.  These  fragile  flies  are  the  favourite  prey  of 
the  endemic  predaceous  wasps  of  the  family  Mimesidae,  and  some  of  the  Crabronidae  like- 
wise gather  them.  Consequently  one  may  find  the  females  of  these  wasps  investigating 
damp,  dark  places,  where  the  Dicranomyia  are  likely  to  be  found,  but  which  ordinarily 
would  have  but  little  attraction  for  such  sun-loving  insects. 

Writing  of  the  Crabronidae,  Perkins  says  (page  Ixxxvi  of  the  reference 
cited) : 

Xenocrabro  hawaiiensis  on  one  occasion  was  caught  carrying  off  Lispe  [Anthomyiidae] 
and  on  another  occasion  a  species  of  the  limnobiid  Dicranomyia. 

De  Meijere  (1920:59)  records  Crabro  (Rhopalum)  tibiale  as  provisioning 
its  nest  principally  with  Molophilus  armatus  Meij. 

Parasitic  natural  enemies 
External  parasites 

Besides  the  predatory  forms  just  considered,  the  various  stages  of  the 
existence  of  many  crane-flies  are  threatened  with  parasites  which  are 
equally  effective  altho  more  insidious  in  their  method  of  attack.  There 
are  unquestionably  many  parasites  of  crane-flies  concerning  which  nothing 
at  all  is  known  at  this  time.  It  is  inconceivable  that  such  species  as 
Holorusia  grandis,  Tipula  abdominalis,  and  other  large  and  common  forms 
do  not  serve  as  hosts  for  parasites  as  yet  unknown.  It  is  this  field  of 
investigation  that  now  promises  some  of  the  most  valuable  results. 

Hexapoda, —  The  only  parasitic  insects  of  which  the  writer  knows 
are  the  dipterous  Tachinidae.  Glover  (1874)  states  that  in  Europe  certain 
Tipulae  are  destroyed  by  a  proctotrupid  parasite  of  the  genus  Diapria 
Latreille.  Kieffer,  in  his  monograph  of  the  Diapriidae  (1911),  does  not 
mention  this  and  the  writer  has  not  seen  it  referred  to  elsewhere.  It  is 
probable  that  the  reference  pertains  to  some  of  the  smaller  Tipula-like 
forms  of  another  family.  However,  it  is  very  strange  that  one  or  more 
of  the  many  families  of  parasitic  Hymenoptera  do  not  infest  some  stage 
of  the  Tipulidae,  and  it  may  be  confidently  predicted  that  such  parasites 
will  one  day  be  discovered. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  731 

'Tachinidae:  Most  of  the  species  of  the  interesting  genus  Siphona 
(Bucentes)  are  parasitic  on  the  caterpillars  of  various  Lepidoptera.  Two 
have  been  recorded  as  parasites  of  the  larvae  of  species  of  Tipula.  The 
better-known  of  these  is  Siphona  cristata  (Fabr.),  a  constant  parasite  of 
the  larvae  of  Tipula  maxima  (Beling  1886,  Czizek  1913,  Kiedel  1913, 
Thompson  1915).  The  method  of  attack  by  the  parasite  has  been 
described  by  Roubaud  (1906).  The  large,  massive  larvae  of  this  species 
of  crane-fly  are  amphibious  or  semi-aquatic,  living  near  the  margins  of 
streams  and  other  bodies  of  water.  They  live  in  the  mud  or  in  the  water 
beneath  the  surface,  at  times  coming  up  to  breathe  the  air  thru  the  two 
large  spiracles  at  the  posterior  end  of  the  body.  It  is  at  this  time,  when 
the  spiracular  disk  is  projected  thru  the  surface  film  and  the  spiracles  are 
open,  that  the  tachinid  is  supposed  to  dart  down  and  deposit  its  eggs  in 
the  open  stigmata.  The  young  parasite  is  somewhat  elongated,  yellow- 
ish orange  in  color,  and  about  a  millimeter  in  length,  and  is  completely 
inclosed  in  a  firm,  membranous  cyst.  It  is  connected  with  one  of  the 
two  principal  tracheal  trunks  of  the  host  by  a  sort  of  chitinized  calyx 
which  opens  into  the  trachea.  At  the  time  of  their  pupation  the  para- 
sites detach  their  organs  of  fixation,  and  with  their  heads  perforate  the 
skin  of  the  host  and  enter  the  humic  earth.  In  nature  this  departure 
coincides  almost  exactly  with  the  time  of  leaving  the  water  by  the  host 
for  the  purpose  of  pupation  in  the  earth.  The  pupal  duration  of  the  para- 
site is  about  three  weeks.  Rennie  (1912)  gives  Siphona  geniculata  (De 
Geer)  as  a  parasite  of  an  undetermined  species  of  Tipula. 

Species  of  the  genus  Admontia  are  important  parasites  of  the  larvae  of 
crane-flies.  In  Europe,  Admontia  arnica  (Meig.)  is  parasitic  on  species  of 
Tipula,  while  in  America  A.  pergandei  Coq.  is  an  important  parasite  of 
the  smoky  crane-fly,  Tipula  cunctans  (Hyslop,  1910:128).  In  the  collec- 
tion of  the  Illinois  Natural  History  Survey  are  several  specimens  of 
A.  pergandei  bred  from  larvae  of  the  smoky  crane-fly  (larvae  taken  at 
Urbana,  Illinois,  parasites  emerged  October  9  to  25, 1900).  Averin  (1913) 
and  Lutchnik  (1916),  in  Russia,  record  the  larvae  of  a  Tipula,  in  one  case 
at  least  —  that  of  the  economic  species  T.  oleracea  • —  as  being  very  heavily 
infested  by  an  undetermined  tachinid  which  may  have  been  the  Admontia 
species  mentioned  above. 

A  third  genus  of  Tachinidae,  Sturmia,  has  recently  been  reported  to 
the  writer  by  J.  D.  Tothill  as  being  parasitic  on  the  larva  of  a  large  crane- 


732  CHARLES  PAUL  ALEXANDER 

fly,  undoubtedly  a  tipuline  form.  The  larva  was  found  at  Chelsea,  near 
Ottawa,  on  May  27,  1906,  by  Dr.  James  Fletcher,  the  parasite  issuing  on 
June  27.  The  tachinid  is  to  be  described  as  Sturmia  tipulensis  Tothill. 

The  genus  Trichoparia  is  parasitic  on  Ctenophora  and  its  allies. 

Altho  the  Tachinidae  are  the  only  dipterous  parasites  that  are  definitely 
known  from  the  Tipulidae,  the  writer  has  a  record  of  a  species  of  Phaonia, 
of  the  family  Anthomyiidae,  which  possibly  is  a  parasite.  Among  some 
Diptera  sent  to  Malloch  for  naming  was  one  specimen  of  a  Phaonia  bearing 
the  label  "  Bred  from  tipulid  pupa,  ace.  no.  14022."  The  writer  is 
indebted  to  R.  H.  VanZwaluwenburg  for  the  following  facts  concerning 
this  specimen:  The  material  was  taken  by  Harry  L.  Parker  on  June  6, 
1916,  on  South  Mountain,  near  Hagerstown,  Maryland.  Under  litter 
and  growth  of  Virginia  creeper  on  rock  a  tipulid  pupa  was  found.  The 
pupa  had  been  eaten  out  inside  and  there  was  a  hole  in  the  side  of  its  body. 
About  a  half  inch  away  from  this  pupa  was  found  a  newly  formed  pupa 
of  an  anthomyiid  fly.  The  adult  emerged  on  June  23,  1916,  and  was 
pinned.  It  is  possible,  of  course,  that  the  Phaonia  was  merely  feeding 
on  the  tipulid,  as  recorded  for  this  genus  elsewhere  in  this  paper  (page 
728),  and  that  the  species  is  a  predatory  enemy  rather  than  a  parasite. 

Arachnida. —  Young  mites  of  species  of  Trombidium,  Rhyncholophus, 
and  other  genera  of  Arachnida,  are  frequently  found  attached  to  the  body 
and  the  wings  of  adult  crane-flies.  This  attachment  is  by  anal  filaments, 
and  it  is  doubtful  whether  any  real  injury  results  to  the  host  therefrom. 
These  cases  are  probably  instances  of  phoresy  to  provide  a  means  of  dis- 
persal of  the  mite.  At  various  times  the  writer  has  found  in  northeastern 
America  a  large  number  of  adult  crane-flies  so  infested,  and  practically 
every  genus  and  many  species  are  included  among  the  number. 

Fungi. —  Among  the  enemies  of  crane-flies,  both  to  the  immature  and 
to  the  adult  stages,  probably  none  are  more  lethal  in  their  action  than 
certain  parasitic  fungi.  The  largest  groups  of  insect-fungous  parasites 
are  the  Entomophthoraceae  (Phycomycetes)  and  the  Laboulbeniaceae 
( Ascomy cetes) .  Dr.  Roland  Thaxter  informs  the  writer  that,  altho  the 
latter  group  infest  other  Diptera,  he  has  never  seen  specimens  on  Tipulidae, 
and  none  have  been  recorded  in  the  literature.  The  writer  is  greatly 
indebted  to  Dr.  Thaxter  and  to  A.  T.  Speare,  Government  Mycoentomolo- 
gist,  for  data  on  this  subject.  Dr.  Thaxter  sent  to  the  writer  for  deter- 
mination a  considerable  number  of  specimens  of  crane-flies  with  their  para- 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  733 

sites,  these  including  the  types  of  many  of  his  Entomophthoraceae  and 
some  new  species  not  yet  described.  *  From  these  data  and  the  literature 
the  following  notes  may  be  given: 

Class,  Phycomycetes 
Family,  Entomophthoraceae 

I.  Entomophthora  (Empusa)  pachyrrhinae  Arthur.     On  the  larvae  of  Nephrotoma  ferruginea-, 
a  manuscript  name. 

2    E.  (E.)  sepulchralis  (Thaxt.).     On  an  adult  Tipula  of  the  subumcolores  group. 

3.  E.  (E.}  caroliniana  (Thaxt.)  [=  E.  arrenoctona  Giard].     On  an  adult   male  of    Tipula 
entomophthorae. 

4.  E.  (E.)  arrenoctona  Giard.     On  male  adults  of  Tipula  paludosa  (Giard  1888,  Villeneuve 
'  1910,  Picard  1913). 

5.  E.  (E.)  tipulae  Fres.     On  Tipula  maxima  (Lohde,  1872). 

6.  E.  (E.}  conglomerata  (Sorokin).     A  species  which  is  probably  this  one  is  recorded  by 
Thaxter  on  larvae  and  adults  of  Tipulidae. 

7.  E.  (E.)  sphaerosperma  Fres.     Recorded  by  Thaxter  from  adult  Tipulidae. 

8.  E.  (E.}  grylli  Fres.     Recorded  on  at  least  two  occasions  from  the  adult  flies  of  Tipula 
ultima  Alex.  [=  T.  flavicans  Fabr.]. 

9.  E.  (E.}  punctata  Thaxt.,  ms.     On  adult  females  of  Pseudolimnophila  noveboracensis. 

10.  E.  (E.}  dipterigena  (Thaxt.).     On  the  adult  flies  of  several  genera  of  Limnobiinae,  as 
'  follows:     Molophilus    hirtipennis,   Limnophila   aprilina,    Pseudolimnophila   nigripleura, 

Ulomorpha  pilosella,  Penthoptera  albitarsis. 

II.  Isaria  sp.     On  adults  of   Dicranomyia  pubipennis.     Additional  undetermined  species 
of  this  same  genus  on  various  tropical  Tipulidae. 

Class,  Hyphomycetes. 
Family,  Mucediniaceae. 

12    Sporotrichum  densum  Link.     On    Tipula  cunclans  Say  [=  T.  infuscata  Lw.J   (Hyslop, 
1910:130). 

Internal  parasites 

Crane-fly  larvae  and  adults  are  infested  by  numerous  protozoan 
and  bacterial  parasites,  the  more  conspicuous  and  constant  being  the 
Sporozoa  (Gregarinidae)  and  the  Bacteria.  General  references  to  this 
subject  may  be  found  in  Leger  (1892),  Labbe  (1899),  and  Minchin 
(1903).  More  especial  references  are  made  to  the  parasites  of  Ptychoptera 
by  Leger  and  Duboscq  (1909),  to  those  of  Dicranota  by  Miall  (1893: 
237),  to  those  of  Ctenophora  by  Anthon  (1908:542),  and  to  those  of 
Tipula  by  Mackinnon  (1913).  In  the  case  of  Ctenophora  the  parasites 
were  frequent  between  the  cells  of  the  alimentary  canal,  especially  in 
the  proventricular  caeca.  The  writer  has  found  a  large  gregarine  very 
abundant  in  the  alimentary  canal  of  the  larva  of  Pedicia  albivitta, 
many  of  the  individuals  being  very  heavily  infested.  Gamkrelidze 
(1913  b)  records  gregarine  and  nematode  parasites  in  Gnophomyia 
larvae. 


734  CHARLES  PAUL  ALEXANDER 

The  more  important  recorded  protozoan  parasites  are  as  follows: 

Class,  Sporozoa. 
Subclass,  Telosporidia. 
Order,  Coccidiomorpha. 
Family,  Adeleidae. 

1.  Adelea  tipidae  Leger.     In  the  intestine  of  species  of  Tipula. 
Order,  Gregarinida. 

Family,  Gregarinidae. 

2.  Hirmocystis  polymorpha  (Le"ger,  1892: 113).     In  the  intestine  of  the  larva  of  Limnobia  sp. 

3.  H.  ventricosa  (Leger,  1892  :111).     In  the  alimentary  canal  of  Tipula  oleracea,  Nephrotoma 
pratensis,  and  other  species. 

4.  Gregarina  longa  (Leger,  1892:117).     In  the  alimentary  canal  of  a  species  of  Tipula. 
Family,  Actinocephalidae. 

5.  Actinocephalus  tipulae  Leger   (1892:141).     In  the  alimentary  canal  of  Tipula  larvae. 
Probably  the  same  species  has  been  recorded  from  the  larvae  of  a  species  of  Ctenophora. 

6.  Pileocephalus  striatus  Leger  &  Duboscq  (1909:887-893).     In  the  mid-intestinal  epithelium 
of  the  larva  of  Ptychoptera  contaminate,.     The  Pileocephalus  live  in  the  epithelium  of  the 
mid-intestine,  attaching  themselves  to  the  epithelial  cells  and  hypertrophying  the  adjoin- 
ing tissues.     They  obtain  their  nutriment  from  the  food  that  penetrates  into  the  cells. 

Family,  Stylorhynchidae. 

7.  Near  Stylorhynchus  (Miall,  1893:237).     In  the  stomach  of  larvae  of  Dicranota  bimaculata. 

Subclass,  Neosporidia. 
Order,  Cnidosporidia. 
Suborder,  Microsporidia. 
Family,  Nosematidae. 

8.  Nosema  strictum  Monz.  (Moniez,  1887).     In  muscles,  conjunctive  tissue,  and  other  parts 
of  Nephrotoma  pratensis. 

9.  Gurleya  francottei  Le"ger  &  Duboscq  (1909:894).     In  the  epithelium  of  the  mid-intestine 
of  the  larva  of  Ptychoptera  contaminata. 

Class,  Flagellata. 

Family,  Trypanosomidae  (Herpetomonadidae). 

10.  Crithidia  campanulata  Leger.     At  the  juncture  of  the  mid-  and  hind-intestines  in  the 
larva  of  Ptychoptera  contaminata  (Leger  &  Duboscq,  1909:898-900). 

The  writer  is  indebted  to  Dr.  R.  Kudo  for  assistance  in  determining  the 
terminology  used  above. 

Bacteria  are  frequently  found  in  crane-fly  larvae.  Leger  and  Duboscq 
(1909:900-901)  record  undetermined  spirochaetes  in  the  epithelial  cells 
of  the  posterior  part  of  the  mid-intestine  of  Ptychoptera  contaminata. 
Dr.  Hugh  Glasgow,  of  the  Geneva  Experiment  Station,  informs  the  writer 
that  in  Illinois  a  large  tipulid  larva,  probably  that  of  Tipula  abdominalis, 
living  in  the  leaf-drift  of  prairie  streams,  is  heavily  infested  with  bacteria. 
Most  of  the  specimens  observed  had  an  abundance  of  small  coccus  and 
spirochaete  forms,  with  occasional  specimens  of  a  gigantic  bacillus  measur- 
ing from  forty  to  eighty  microns  in  length  and  disporous.  These  large 
bacilli  infest  the  hind-gut  of  the  larva. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  735 

HISTORICAL    SUMMARY    OF    THE    IMMATURE    STAGES 

The  discovery  and  description  of  the  immature  stages  of  crane-flies 
has  by  no  means  kept  pace  with  the  taxonomic  work  that  has  been  done 
on  the  adult  flies.  There  are  a  very  large  number  of  common  species  in 
every  country  concerning  whose  early  stages  nothing  at  all  is  known, 
and  the  knowledge  that  exists  concerning  a  comparatively  small  number 
results  from  the  labors  of  a  few  students.  Among  these  may  be  mentioned 
Beling,  Mik,  Brauer,  Hart,  Needham,  and  Malloch.  Some  other  workers 
have  given  excellent  descriptions  of  single  species,  while  still  others  have 
contributed  important  treatises  on  the  anatomy,  morphology,  histology, 
and  related  subjects  of  the  different  species.  The  descriptions  of  the 
earlier  workers  are,  for  the  most  part,  of  historical  interest  only,  but 
in  a  few  cases  they  are  exceptional  and  are  still  the  main  source  of  informa- 
tion concerning  certain  species.  Many  of  the  foremost  workers  on  the 
taxonomy  of  the  adult  flies  have  summarized  the  earlier  writings  on  the 
immature  stages  but  have  themselves  contributed  little  to  the  subject, 
among  these  being  Schiner,  Loew,  Osten  Sacken,  Skuse,  and  Brunetti. 
The  following  chronological  summary  gives  an  indication  of  the  more 
important  work  that  has  been  done  on  this  phase  of  the  subject: 

1722 Frisch Tipula  oleracea 

1740 De  Reaumur Ptychoptera,  Tipula 

1776 De  Geer Phalacrocera,  Ctenophora,  Tipula 

1803 Schellenberg Cylindrotoma 

1829 Stannius 'Limnobia  xanthoptera  [=bifasciata] 

1832  (Posthumous).  Lyonet Ptychoptera 

1833 Bouche 'Dictenidia  sp.,  Tipula  sp. 

1834 Von  Roser Hexatoma  nigra 

1838 Boie Cylindrotoma 

1840 Dufour Trichocera 

1842 Zeller Cylindrotoma 

1846 Bremi-Wolf Trichocera,  Limnobia    xanthoptera  [=bifasciata], 

Nephrotoma 

1849 Perris Trichocera,  Limnophila  dispar,  Ula,  Tanyptera 

1854 Brauer,     Egger,    and 

Frauenfeld Chionea 

1867 Nowicki Dactylolabis  wodzickii 

1872 Weyenbergh Ctenophora,  Dictenidia:  anatomy  and  histology 

1873-86 Beling The  immature  stages  of  69  species  of  European 

Tipulidae,  including  30  species  of  Tipula 

1875-76 Hammond Tipula  oleracea:  anatomy 

1876 Grobben Ptychoptera:  anatomy 

1876-1902 De  Rossi Liogma  glabrata,  Tanyptera  atrata 

1880 Hermann Tanyptera  atrata 

1882-1900 Mik Discobola,  Elliptera,  Dactylolabis,  Tipula  rufina, 

etc. 


736  CHARLES  PAUL  ALEXANDER 

1883 Brauer Important    work   on    the    classification    of    the 

immature  stages 

1884 Gercke Rhamphidia  longirostris,  Tanyptera  atrata 

1890 Van  Gehuchten Ptychoptera :  histology  of  alimentary  canal 

1893-97 Miall Dicranota,  Phalacrocera 

1895  [1898] Hart Pioneer  American  worker;  Bittacomorpha,  etc. 

1897 Bengtsson Phalacrocera :  anatomy 

190Q-09 Johnson Teucholabis,  Aeshnasoma,  etc. 

1901 Kellogg Holorusia  grandis  (as  rubiginosa) :  anatomy 

1901 Miiggenberg Liogma  glabrata 

1901-08 Needham Dicranomyia  simulans,  Pedicia  albivitta,  Rhaphi- 

dolabis,  Tipula  ultima  (as  flavicans) 

1907-08 Steinmann Triogma  trisulcata 

1908 Holmgren Phalacrocera:  mouth  parts 

1908 Anthon Ctenophora  angustipennis :  anatomy 

1908-09 Pastejrlk Limnobia  xanihoptera,  Ctenophora 

1908-09 Mtiller Triogma  trisulcata 

1909 Brocher Tipula  lunata 

1909 Greene Tipula  trivittata 

1909 Thienemann Dicranomyia  trinotata 

1910 Brown Tipula  maxima :  anatomy 

1910 Hyslop Tipula  cunctans  (as  infuscata) 

1911. Vimmer Ctenophora,  Tipula  oleracea:  mouth  parts 

1911-16 De  Meijere Trentepohlia   pennipes,    Tipulodina   pedata,   and 

several  European  species 

1912 Keilin Trichocera:  morphology 

1913 Caudell Tipula  ultima  (as  flavicans) 

1913 Gerbig Anatomy   of   the   respiratory   system   of   many 

European  species  of  Tipulidae 

1913 Picado Trentepohlia  bromeliadicola 

1914 Del  Guercio Tipula  oleracea 

1914^16 Topsent Ptychoptera  albimana 

1914-19 Alexander Dicranoptycha,  L-la,  Eriocera,  Hexatoma,  Penth- 

optera,  Liogma,  Prionocera,  Tipula  arctica 

1915 Swezey Dicranomyia  foliocuniculator 

1915 Lovett Ctenophora  angustipennis 

1915 Wesenberg-Lund Dicranota,  Phalacrocera,  Triogma 

1915-17 Malloch Limnobia  immatura,   Gnophomyia,   and,   in   his 

uncertain  material,  Antocha,  Elephantomyia 

1916-17 Rennie Tipula  paludosa 

1918 Cameron Cylindrotoma  splendens 

1920 Hudson Gnophomyia  rufa,  Limnophila  sirristra 

1920 Lenz Thaumastoptera,  Cylindrotominae 

ECONOMIC    IMPORTANCE 

The  larvae  of  some  species  of  crane-flies,  almost  all  belonging  to  the 
genera  Tipula  and  Nephrotoma,  often  do  considerable  damage  to  various 
crop  species,  the  injury  being  largely  caused  by  the  larvae  devouring  the 
roots  and  thus  killing  the  plants.  Certain  of  these  crane-fly  species,  as 
Tipula  oleracea,  T.  paludosa,  and  Nephrotoma  maculata  in  Europe,  and 
Tipula  bicornis  and  Nephrotoma  ferruginea  in  America,  have  long  been 
known,  but  several  others  have  come  into  prominence  only  within  the 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  737 

past  ten  years,  notably  '  Tipula  parva  Lw.  (supposition)  in  Japan,  and 
T.  cunctans  and  T.  simplex  in  North  America.  The  more  important  out- 
breaks of  this  nature  which  have  been  recorded  are  as  follows. 

Species  affecting  herbaceous  plants 

Nephrotomaferruginea  has  been  reported  by  Webster  (1891,  and  1893  a: 
243-245)  as  injuring  young  wheat  in  Indiana,  the  injury  in  some  cases 
being  estimated  at  fifty  per  cent. 

Tipula  oleracea  is  the  best-known  European  species  in  this  group,  infest- 
ing a  wide  range  of  plants.  The  more  important  outbreaks  of  this  species 
as  recorded  are  as  follows:  Ewert  (1899)  reports  from  Germany  an 
unusually  severe  attack,  in  which  the  larvae  of  this  species  and  of  Tipula 
nigra  did  a  considerable  amount  of  damage  to  the  roots  of  grasses.  They 
were  so  abundant  that  from  ten  to  twenty  could  be  found  in  the  area  of 
one  square  foot.  Grasses  in  the  infested  area  were  completely  destroyed. 
Ormerod  (1885, 1886, 1900)  records  serious  injury  by  T.  oleracea  to  pastures 
and  meadows  in  England,  and  recommends  applications  of  guano  either 
alone  or  mixed  with  salt,  kainit,  or  superphosphate.  This  species  had  been 
recorded  also  as  being  a  serious  pest  in  Ireland  (Anonymous  reference, 
1904b)  in  grainfields  and  meadows,  and  careful  plowing  and  the  use  of 
fertilizers  for  its  control  are  advised.  Schoyen  (1903),  in  Norway,  reports 
serious  injury  to  meadows  and  pastures,  as  well  as  to  cereals  and  young 
cabbage  plants.  Del  Guercio  (1914)  records  serious  injury  in  the  Italian 
rice  fields. 

A  few  additional  outbreaks  of  Tipula  oleracea  may  be  mentioned. 
Barthou  (1913)  records  injury  to  canes  (raspberry,  strawberry,  and 
others)  in  France.  Desoil  (1914)  reports  injury  to  meadows  in  France, 
and  Ritzema  Bos  (1915)  to  meadows  in  Holland. 

Tipula  paludosa  is  reported  by  Lind,  Rostrup,  and  Kolpin  Ravn  (1914 
and  1915)  as  causing  serious  injury  to  oats  and  barley  in  Denmark,  and 
Rennie  (1916  and  1917)  reports  the  same  species  as  injuring  corn  and 
pastures  in  England. 

Tipula  flavolineata  is  reported  by  Sopotzko  (1916)  as  injuring  clover 
in  Russia.  The  specific  identity,  however,  is  probably  erroneous,  as  this 
species  lives  almost  entirely  in  .decaying  wood  (Beling,  1873  b:  581-582). 

Onuki  (1905)  records  Tipula  parva  Lw.  (supposition)  as  one  of  the  seri- 
ous rice  pests  in  Japan.  In  some  localities  from  sixty  to  ninety  per 


738  CHARLES  PAUL  ALEXANDER 

cent  of  the  seedlings  have  been  destroyed  by  this  species.  The  larvae 
apparently  cannot  exist  in  water  for  any  long  period  of  time,  and  so  may 
be  killed  by  flooding  the  fields  for  from  six  to  thirty-six  hours.  Del 
Guercio  (1914)  offers  the  same  recommendation  for  the  control  of  T. 
oleracea  in  the  Italian  rice  fields. 

Tipula  bicornis  has  been  found  in  grass  and  clover  lands  in  Illinois  by 
Forbes  (1890),  and  on  clover  in  Indiana  by  Webster  (1892 a). 

Tipula  cunctans,  as  recorded  by  Hyslop  (1910:126,  as  T.  infuscata), 
works  largely  on  Japan  clover  and  other  leguminous  plants.  The  larvae 
are  often  exceedingly  numerous,  as  many  as  two  hundred  having  been 
found  in  an  area  covering  but  little  more  than  one  square  foot  of  soil. 
They  destroy  the  plants  by  devouring  the  roots  and  sucking  the  juices. 
It  is  recommended  that  the  infested  fields  be  plowed  under  before  the 
adult  flies  emerge  (in  the  autumn),  and  the  following  season  be  planted 
to  corn  or  potatoes  or  else  left  fallow. 

Tipula  simplex  has  been  found  on  pasture  land  and  alfalfa  in  Cali- 
fornia by  Doane  (1908)  and  by  Carnes  and  Newcomer  (1912).  Doane 
(cited  by  Hyslop,  1910:120-121)  also  reports  an  outbreak  of  this  species 
on  wheat  and  grass  lands  and  in  clover  fields  in  central  California  in 
1907,  and  states  that  thousands  of  acres  of  these  crops  were  stripped  of 
their  verdure. 

The  following  summary  gives  the  various  species  of  plants  that  are 
damaged  or  destroyed  by  crane-flies  in  general.  Very  many  of  the 
injuries  reported  for  Tipula  oleracea,  however,  are  omitted. 

Family  Crop  Species  Authority 

Gramineae Wheat Nephrotoma  ferruginea  Webster  (Indiana),  1891, 1893 a 

Wheat Tipula  simplex Hyslop  (California),  1910 

Wheat Tipulidae Stedman  (Missouri),  1902 

Corn Tipula  paludosa Rennie  (England),  1917 

Cora Tipula  sp Kirk  (New  Zealand),  1895 

Barley Tipula  paludosa Lind,  Rostrup,  and  Kolpin  Ravn 

(Denmark),  1915 
Barley Tipula  sp Wahl    and    Miiller    (Germany) , 

1914 

Barley Tipula  oleracea Goriatchkovsky  (Russia),  1915 

Oats Tipula  paludosa Lind,  Rostrup,  and  Kolpin  Ravn 

(Denmark),  1914,  1915 

Rice Tipula  oleracea Del  Guercio  (Italy),  1914 

Rice Tipula  parva Onuki  (Japan),  1905 

Cereals Tipula  oleracea Anonymous  reference  (Ireland) , 

1904  b 
Cereals Nephrotoma  pratensis, ,  Hollrung  (Germany),  1898 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II 


739 


Family 
Gramineae  .  . 


Crop  Species 

Meadows     and 

pastures Tipula  bicornis . 

Meadows      and 

pastures Tipula  simplex . 


Meadows      and 

pastures Tipula  oleracea 

Meadows  and 

pastures Tipula  oleracea 

Meadows  and 

pastures Tipula  paludosa 

Meadows  and 

pastures Tipula  sp 

Meadows  and 

pastures Tipula  sp 

Meadows  and 

pastures Tipula  sp 


Meadows      and 

pastures Tipula  oleracea 


Liliaceae  , 

,  .  .   Tulips  (bulbs). 

Nephrotoma  maculata  . 

Cruciferae.  .  .  . 

Polygonaceae.  , 
Rosaceae 

Onions  and  gar- 
lic   
.  .   Cabbage  
Cabbage  
Cabbage  
Cabbage  
Cabbage  
.  .   Buckwheat  .... 
Roses 

Tipula  oleracea  
Tipula  oleracea  
Tipula  oleracea  
Tipula  oleracea  
Tipula  sp  
Nephrotoma  sp  
Tipula  oleracea  
Nephrotoma  lineata 

Leguminosae  .  , 

Raspberry    and 
strawberry.  .  . 
.  .   Clover  

Tipula  oleracea  
Tipula  bicornis  

Clover  

Tipula  bicornis  

Clover 

Tipula  flavolineata 

Clover  
Japan  clover  .  .  . 

Tipula  oleracea  
Tipula  cunctans  

Alfalfa.. 

Tipula  simplex  .  . 

Peas Tipula  paludosa 


llmbellif  erae . 
Solanaceae . . 


Compositae . 


Carrots Tipulidae 

Potatoes Tipula  oleracea 

Potatoes Tipula  lateralis 

Potatoes,  stored  Trichocera 

Potatoes,  stored  Trichocera 

Cardoon Tipula  sp 


Authority 
Forbes  (Illinois),  1890 

Games    and    Newcomer    (Cali- 
fornia), 1912 

Anonymous  reference   (Ireland), 

1904  b 
Desoil  (France),  1914 

Rennie  (England),  1917 
MacDougall  (Scotland),  1915 
Ritzema  Bos  (Holland),  1915 

Marchal  and  Prillieux  (France), 
1916 

Ormerod  (England),  1885,  1886, 

1900 
Collinge  (England),  1911 

Vassiliev  (Russia),  1915 
Schoyen  (Norway),  1903 
Averin  (Russia),  1913 
Goriatchkovsky  (Russia),   1915 
Lutchnik  (Russia),  1916 
Lutchnik  (Russia),  1916 
Goriatchkovsky  (Russia),  1915 
Schaufuss  (Germany),  1901 

Barthou  (France),  1913 
Webster  (Indiana),  1892 a 
Forbes  (Illinois),  1890 
Sopotzko  (Russia),  1916 
Del  Guercio  (Italy),  1914 
Hyslop  (Tennessee),  1910 
Games    and    Newcomer     (Cali- 
fornia), 1912 
Lind,  Rostrup,  and  Kolpin  Ravn 

(Denmark),  1914 
Lesne  (France),  1905 
Beling  (Germany),  1887 
Cameron  (Scotland),  1917 
.Johannsen  (Maine),  1910 
Carpenter  (Ireland),  1912 
Lesne  (France),  1908 


Species  affecting  woody  plants 

Species  that  injure  or  destroy  living  trees  or  shrubs  are  not  common, 
and  the  damage  that  they  do  is  almost  entirely  of  a  minor  nature. 


740  CHARLES  PAUL  ALEXANDER 

Ctenophora  angustipennis  is  recorded  as  doing  secondary  damage  to  prune 
trees  in  Oregon  (Lovett,  1915).  Fuchs  (1900)  records  four  species  — 
Tipula  scripta,  T.  marginata,  Nephrotoma  cornicina  [as  iridicolor],  and 
N.  quadrifaria  —  as  injuring  young  plants  of  both  deciduous  and  conif- 
erous species.  Taschenberg  (1880:54)  describes  Tipula  flavolineata  and 
Nephrotoma  crocata  as  destroying  year-old  seedlings  of  fir  and  larch. 
Matsumura  (1916:466)  records  the  larva  of  Nephrotoma  makiella  as 
injurious  to  the  mulberry  (Morus)  in  Formosa. 

METHODS    OF   EXPERIMENTAL    PROCEDURE 

Collecting 

The  larvae  of  the  larger  species  of  crane-flies,  such  as  those  of  the  genera 
Tipula,  Pedicia,  and  Eriocera,  may  be  readily  discovered  in  the  field, 
but  the  immature  stages  of  the  smaller  Tipulidae  are  not  so  easily  found. 
It  is  necessary  to  bring  into  the  laboratory  large  quantities  of  the  material 
in  which  the  immature  stages  are  supposed  to  be  living,  and  there  to 
examine  it  with  considerable  care.  In  the  case  of  mud  or  earth,  it  is 
better  to  wash  away  the  finer  silt  particles  and  examine  the  residue. 
The  writer  has  found  it  most  convenient  to  use  a  Simplex  water-net, 
placing  in  it  a  handful  of  earth  and  holding  it  underneath  a  running  faucet . 
The  mesh  of  this  net  is  of  sufficient  diameter  to  retain  any  crane-fly  larvae 
except  the  very  young  stages.  When  the  fine  particles  have  been  removed 
the  residue  can  be  examined  in  water  in  white  enameled  or  porcelain 
dishes,  and  the  larvae  and  pupae  may  be  easily  detected  in  the  water. 
Numerous  associated  forms  of  life  will  be  found,  and  these  should  be  pre- 
served or  recorded. 

The  immature  stages  of  wood-inhabiting  species  are  most  easily  found 
in  the  field  by  a  patient  and  painstaking  examination  of  the  removed 
bark  and  the  exposed  parts  of  the  tree  stump  or  trunk.  Moss-inhabit- 
ing species,  such  as  those  of  Liogma  and  many  tipuline  forms,  may  be 
shaken  from  their  haunts  onto  a  piece  of  white  oilcloth,  where  they  are 
easily  observed. 

Killing  and  preserving 

The  larvae  and  pupae  to  be  studied  should  be  dropped  into  boiling  water 
for  a  brief  instant.  The  larva,  on  contact  with  the  water,  at  once  expands 
to  its  maximum  size,  the  spiracular  disk  is  spread  wide  open,  and  the  anal 


THE  CRANE-FLIES  or  NEW  YORK  —  PART  II  741 

gills  are  completely  everted.  The  specimens  should  be  placed  in  4-per- 
cent formalin  or,  preferably,  70-  to  75-per-cent  alcohol.  Large,  fleshy 
larvae,  such  as  those  of  the  Tipulinae,  should  be  slit  at  the  third  or  the 
fourth  abdominal  segment  to  allow  easy  penetration  of  the  preservative. 
Other  notes  on  preserving  material  are  given  by  Banks  (1909)  and  by 
Williamson  (1916). 

Study 

In  the  study  of  the  gross  material,  both  the  binocular  and  the  com- 
pound microscope  may  be  used.  A  special  word  on  the  preparation 
of  the  head  capsule  of  the  larva  may  be  helpful.  In  removing  the  head 
capsule  from  the  body  it  is  generally  easiest  to  slit  longitudinally  the  tho- 
racic segments  back  of  the  head  and  pull  the  capsule  thru  this  incision. 
In  forms  with  compact  and  massive  capsules,  the  thorax  and  the  head  may 
be  snipped  off  and  the  head  everted  after  the  manner  of  turning  the 
finger  of  a  glove.  The  capsule  should  be  left  in  a  5-per-cent  solution 
of  caustic  potash  until  all  the  fleshy  parts  have  been  removed.  It 
should  be  dissected  out  so  that  all  the  mouth  parts  are  isolated  and 
rendered  distinct  for  study.  Then  follow  the  usual  processes  of  washing 
and  dehydration,  and  the  final  mounting  in  Canada  balsam. 

Various  means  are  available  for  examining  the  spiracular  disk.  The 
specimen  may  be  placed  in  a  watch  crystal  under  water  and  held  firmly 
in  place  by  a  piece  of  glass,  as,  for  example,  a  broken  microscope  slide. 
By  holding  a  heavy  piece  of  glass  over  the  anterior  end  of  the  body,  the 
posterior  end  may  be  bent  at  a  righ't  angle  and  bolstered  in  place  by  two 
or  three  other  glass  fragments  so  that  the  disk  is  directed  straight  upward 
and  its  details  are  easily  examined.  Other  methods,  such  as  embedding 
the  anterior  end  of  the  larva  in  paraffin  in  a  deep  dish  and  submerging 
the  body,  may  be  followed.  It  must  be  emphasized  that  in  alcoholic 
gross  material  the  study  should  be  conducted  under  alcohol  or  water. 

The  measurements  of  the  pupa  —  abbreviated  in  the  text  as  d.-s. 
(dextro-sinistral)  for  the  width  and  d.-v.  (dorso-ventral)  for  the  depth  — 
are  taken  opposite  the  wing  pad. 

Rearing 

In  order  to  rear  the  immature  stages  of  crane-flies  successfully  the 
natural  haunts  should  be  imitated  as  closely  as  possible.  In  the  case 
of  wood-inhabiting  species,  large  pieces  of  the  material  in  which  the  larvae 


742  CHARLES  PAUL  ALEXANDER 

are  working  may  be  brought  into  the  laboratory  and  placed  in  the  breed- 
ing cages.  This  method  may  be  followed  with  species  of  Tanyptera, 
Ctenophora,  and  other  genera. 

Fungicolous  species,  as  a  rule,  also  are  easily  reared.  The  entire  fungus 
in  which  the  specimen  is  found  should  be  brought  into  the  laboratory 
and  placed  in  a  jar  on  a  bed  of  clean  sand.  The  sand  takes  up  the  liquids 
produced  by  the  disintegration  of  the  fungus  and  provides  a  place  for 
pupation.  Species  of  Limnobia,  Ula,  and  other  forms  are  reared  in  this 
manner. 

The  chances  for  error  in  rearing  are  many.  One  must  be  certain  that 
there  are  no  other  larvae  in  the  breeding  jar  with  the  one  that  is  being 
reared;  else  one  of  these  other  larvae  might  transform  and  emerge  first, 
and  the  results  would  be  altogether  misleading.  The  writer  has  had 
this  happen  in  his  breeding  cages,  even  after  the  utmost  care  had  been 
used  to  guard  against  it.  Beling,  the  great  German  student  of  the 
immature  stages  of  crane-flies,  made  a  few  mistakes  in  the  same  way; 
as,  for  example,  in  the  case  of  his  Trimicra,  the  larvae  that  he  describes 
being  pediciine  and  probably  a  species  of  Dicranota  or  the  young  larvas 
of  a  Tricyphona.  What  happened,  presumably,  was  that  Beling  found 
these  pediciine  larvae  and  placed  them  in  rearing;  in  the  same  cage,  bu!^ 
unknown  to  the  breeder,  was  a  larva  of  Trimicra  which  emerged,  and 
naturally  Beling  thought  it  came  from  one  of  the  larvae  that  he  had 
placed  in  rearing.  It  is  usually  easy  to  check  up  such  errors.  Thus, 
the  writer  has  placed  in  rearing  the  larvae  of  Penthoptera  and,  to  his 
surprise,  had  adults  of  Limnophila  adusta  emerge.  Obviously  larvae 
of  L.  adusta  got  into  the  cage  in  spite  of  precautions,  and  emerged  first. 
When  closely  related  species  are  concerned,  however,  it  becomes  a  hard 
matter  to  straighten  the  tangle.  Hence  a  species  cannot  be  reared  too 
many  times,  since  each  rearing  checks  up  the  previous  results. 

The  precaution  to  be  taken  in  the  case  of  mud-inhabiting  or  sand- 
inhabiting  species  is  to  see  that  the  mud  or  sand  is  baked  or  thoroly 
desiccated  in  order  to  destroy  all  life  in  it.  Then  it  may  be  remoistened, 
and  the  larva  or  larvae  chosen  to  rear  may  be  put  into  the  earth  without 
the  chance  that  some  unknown  larva  may  be  lurking  in  the  medium  and 
may  emerge  first,  and  so  bring  about  confusion. 

The  writer  has  found  that  the  most  satisfactory  way  to  rear  small 
tipulid  larvae  found  in  earth  or  sand  is  to  place  a  small  amount  of  baked 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  743 

earth  from  their  haunts  in  a  4-  or  6-dram  shell  vial,  moisten  the  earth, 
and  place  a  thin  layer  of  moss  over  it.  The  vial  should  not  be  corked, 
but  should  be  covered  with  a  piece  of  cheesecloth  held  in  place  by  rubber 
bands.  In  the  case  of  carnivorous  species,  as  the  Hexatomini  and  the 
Pediciini,  individuals  should  be  isolated  in  vials,  but  the  herbivorous 
species  may  be  bred  in  large  numbers  in  single  containers.  The  predatory 
forms  are  usually  distinguished  without  difficulty  by  their  extremely 
active,  snakelike  motions,  the  other  species  being  more  sluggish  in  their 
actions. 

Strictly  aquatic  forms,  such  as  Antocha,  are  reared  only  with  great  diffi- 
culty. These  species  can  survive  only  in  rapidly  flowing,  well-aerated 
water,  and  it  is  usually  a  difficult  matter  to  imitate  this  condition  success- 
fully. The  best  plan  is  to  place  rearing  cages  in  the  natural  haunts  of 
the  larvae.  This  can  be  done  if  a  suitable  location  is  sufficiently  close  at 
hand  and  free  from  disturbance  by  inquisitive  passers-by.  The  chief 
source  of  danger  to  breeding  cages  in  natural  streams  is  that  a  sudden 
rise  of  water  may  suffocate  the  larvae  or  wash  the  cages  downstream, 
or  else  may  destroy  the  contents  of  the  cage  by  a  thick  deposition  of  silt. 
Lotic  species  that  are  discovered  in  streams  far  removed  from  the  laboratory 
can  be  transported  alive  for  long  distances  in  folds  of  wet  cheesecloth. 
The  writer  has  found  this  to  be  the  most  satisfactory  way  to  keep  advanced 
pupae  alive  until  they  are  ready  to  emerge  as  adults. 

In  general,  the  writer  has  found  species  of  the  tribe  Pediciini  the  most 
difficult  to  rear,  and  the  Limnobiini  and  the  Eriopterini  perhaps  the 
easiest.  Mud-inhabiting  species  are  easily  reared,  but  species  from  rushing 
torrents  are  at  the  opposite  extreme  and  it  is  almost  an  impossibility  to 
bring  some  of  these  species  thru  to  the  adult  condition. 

TYPES  OF  THE  IMMATURE  STAGES 

The  material  on  which  this  study  is  based  was  almost  entirely  reared. 
It  seems  desirable  that  these  authentic  specimens  of  the  larva  and  the 
pupa  should  be  so  designated  that  they  will  have  a  value  greater  than 
that  possessed  by  ordinary  specimens.  No  terms  for  the  designation  of 
types  of  the  immature  stages  are  known  to  the  writer,  and  the  two  that 
are  needed  in  this  paper  are  here  defined  as  follows: 

Nepionotype,  The  type  of  the  larva. 

Neanotype,  The  type  of  the  pupa. 


744  CHARLES  PAUL  ALEXANDER 

The  type  specimen  should,  of  course,  be  selected  only  when  there  is 
absolute  certainty  of  the  identification,  and  in  most  cases  this  determination 
can  be  made  only  by  rearing  the  species.  After  the  species  has  been  reared 
(this  should  be  done  many  times,  if  possible,  in  order  to  check  up  the 
identity) ,  a  good  representative  specimen  may  be  chosen  as  the  type  of  the 
stage.  In  the  cases  in  which  the  species  is  known  only  from  a  single 
specimen,  the  nepionotype  may  be  the  larval  skin,  the  neanotype  the 
pupal  skin.  The  remaining  specimens  of  the  original  series  become  para- 
types.  The  types  of  the  immature  stages  possess  fully  the  value  of  the 
type  of  the  adult  and  should  be  as  carefully  preserved.  The  types  herein 
designated  are  in  the  collection  of  the  writer.  They  are  preserved  in 
alcohol,  but  the  larval  heads  of  most  species  have  been  removed,  treated 
with  caustic  potash,  and  mounted  in  balsam. 

EXTERNAL  MORPHOLOGY 

The  larvae  and  the  pupae  of  crane-flies  show  considerable  diversity 
in  their  general  form.  The  fundamental  plan  of  structure  remains  much 
the  same  thruout  the  group,  but  the  details  are  widely  different  and  furnish 
the  characters  in  use  for  the  separation  of  the  various  tribes  and  lesser 
divisions. 

The  immature  stages  of  crane-flies  have  evolved  more  rapidly  than 
have  the  adult  flies,  and  in  many  features  they  show  a  greater  specialization. 
The  head  capsule  of  the  larva  seems  to  be  the  most  constant  feature, 
the  same  fundamental  type  of  structure  recurring  in  the  generalized 
members  of  all  the  various  groups,  indicating  a  close  phylogenetic  relation- 
ship. On  the  other  hand,  the  respiratory  organs  of  both  the  larvae  and 
the  pupae  vary  greatly  in  the  different  species  and  are  obviously  molded 
by  habitat.  The  often-repeated  statement  that  the  inside  of  an  organism 
shows  what -it  is,  while  the  outside  shows  where  it  has  been,  is  well  illus- 
trated here. 

The  larva 
General  features 

The  form  of  the  larval  body  is,  as  a  rule,  moderately  elongated  and 
usually  terete.  The  head  is  eucephalous  and  non-retractile  in  the  three 
families  Tanyderidae,  Ptychopteridae,  and  Rhyphidae.  It  is  incomplete 
and  more  or  less  retractile  in  all  the  species  of  Tipulidae.  The  body 
is  shortest  in  the  more  generalized  forms,  becoming  greatly  elongated 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  745 

in  many  Eriopterini  and  some  Limnobiini  (Dicranoptycha) .  It  is  mada 
up  of  the  composite  head  capsule,  three  thoracic  segments,  and  nine 
evident  abdominal  segments.  In  some  species  all  the  abdominal  segments 
are  subdivided,  respectively,  into  a  narrow  basal  and  a  usually  broader 
posterior  ring,  or  annulus;  in  other  species  only  the  basal  segments  are 
so  subdivided.  The  integument  is  usually  covered  with  a  dense  appressed 
pubescence  and  often  bears  setae,  or  pencils  of  hairs,  or,  in  some  Cylin- 
drotominae,  spinous  projections. 

Respiration  is  characteristically  metapneustic ;  in  the  Rhyphidae  it  is 
amphipneustic,  in  Antocha  apneustic.  The  typical  metapneustic  forms 
often  show  vestigial  lateral  spiracles,  but  these  are  not  functional  in  any 
species  known  to  the  writer  and  the  peripneustic  type  of  larva  is  still 
unknown  in  this  group  of  Diptera.  The  spiracles  are  placed  at  the  ends 
of  the  long  breathing  tubes  in  the  Tanyderidae  and  the  Ptychopteridae. 
In  the  Tanyderidae,  the  Tipulidae,  and  the  Rhyphidae  the  disk  is  sur- 
rounded by  a  varying  number  of  lobes  which  are  rarely  indistinct,  these 
ranging  in  number  from  two  to  eight.  Anal  gills  are  found  in -repre- 
sentatives of  almost  all  the  major  groups  of  crane-flies,  and  their  loss  is 
a  result  of  habitat  and  non-usage.  In  wood-inhabiting  species  the  gills 
are  often  modified  into  blunt  lobes,  having  the  evident  function  of  pro- 
pulsion by  shoving. 

Body  form 

As  already  stated,  in  the  majority  of  crane-fly  larvae  the  body  is  terete 
or  approximately  so,  but  in  some  species  it  is  decidedly  depressed  with 
the  ventral  surface  flattened.  Such  forms  are  Dactylolabis,  some  Cylin- 
drotominae,  and  some  Tipulinae.  The  integument  is  produced  into 
elongate  spines  and  blades  in  almost  all  species  of  Cylindrotominae,  similar 
conditions  being  suggested  in  a  few  tipulines.  A  definite  arrangement  of 
setae  (chaetotaxy)  obtains.  The  basal  abdominal  ring  is  provided  with 
a  transverse  creeping- welt  in  the  Limnobiini  and  in  some  Hexatomini 
and  Pediciini,  as  well  as  in  a  few  other  forms.  In  some  genera, 
as  Epiphragma,  this  welt  is  practically  naked;  in  others  it  is  covered  with 
a  microscopic  scurfiness;  while  in  still  others  (Dicranota)  it  is  separated 
into  distinct  paired  prolegs,  which  are  armed  with  circlets  of  chitinized 
hooks  that  lessen  in  size  from  the  tips  basally.  The  welts  are  both  dorsal 
and  ventral  in  position  in  many  Limnobiini  and  in  some  Pediciini  (Rhaphi- 


746  CHARLES  PAUL  ALEXANDER 

dolabina),  or  are  ventral  only  (Rhamphidaria,  Ularia,  Epiphragmaria , 
and  most  Pediciini).  In  number  they  range  from  four  (on  abdominal 
segments  4  to  7)  in  most  Pediciini,  to  five  or  six  (on  abdominal  segments  2 
to  7)  in  Ularia,  Epiphragmaria,  and  other  forms. 

Organs  of  respiration 

The  spiracular  disk. —  The  posterior  end  of  the  body  is  usually  truncated, 
bearing  the  two  spiracles  and  surrounded  by  a  number  of  fleshy  lobes. 
These  lobes  vary  much  in  their  shape  and  armature,  and  range  in  number 
from  two  in  the  Pedicaria  and  the  Antocharia  to  six  or  eight  in  the  Tipulinae. 
The  inner  faces  of  the  lobes  are  often  lined  with  brown  or  black  horny 
plates,  which  serve  as  points  of  attachment  for  the  longitudinal  muscles. 
Their  arrangement  and  distribution  are  of  great  service  in  specific 
classification. 

The  Limnobaria  usually  have  the  lobes  surrounding  the  spiracular 
disk  indistinct;  the  Antocharia  have  two  long  ventral  lobes;  the  Rham- 
phidaria five  lobes;  the  Ellipteraria  and  the  Dicranoptycharia  four  lobes. 

In  the  Eriopterini,  the  Elephantomyaria  and  some  Eriopteraria  have 
four  lobes;  the  majority  of  species  of  the  latter  division  have  five.  In 
these  last-named  species  the  disk  is  almost  squarely  truncated,  and  the 
five  lobes  are  subequal  and  stellate  in  appearance. 

The  Hexatomini  usually  have  four  lobes,  with  the  ventral  pair  longer 
than  the  lateral  pair.  In  a  few  cases  only  (Ularia,  Dicranophragma, 
Poecilostola) ,  a  reduced  mid-dorsal  lobe  is  present.  In  the  more  specialized 
forms  the  lateral  lobes  also  tend  to  reduce,  simultaneously  with  the 
elongation  of  the  ventral  lobes.  In  this  tribe  the  disk  is  fringed  with  long 
hairs,  which  are  sometimes  excessively  elongate,  especially  on  the  ventral 
lobes  where  they  appear  as  long  fringes  of  delicate  hairs  that  spread  out 
on  the  surface  film  of  the  water  as  broad  fans.  Such  fringes  are  found 
in  a  large  number  of  Hexatomini  and  also  in  the  Adelphomyaria  (sup- 
position). Near  the  tips  of  the  ventral  lobes,  but  inside  this  fringe  of 
hairs,  are  often  inserted  one  or  more  stiff  sensory  bristles. 

In  the  Pediciini,  the  Pedicaria  have  two  lobes  and  the  Adelphomyaria 
(supposition)  have  four.  In  the  former  division  the  lobes  are  ventral 
in  position  and  are  almost  naked. 

The  Cylindrotominae  have  six  rather  indistinct  lobes  in  Cylindrotoma, 
and  four  in  the  other  genera. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  747 

The  Tipulinae  show  six  lobes  in  practically  all  genera,  the  only  exceptions 
being  that  there  are  eight  in  a  few  rare  cases  of  Tipula  and  five  in  Doli- 
chopeza,  and  that  lobes  are  indistinct  or  lacking  in  Tanyptera. 

In  the  Rhyphidae  (Trichocerinae)  the  spiracular  disk  is  very  similar 
to  that  in  many  Tipulidae,  and  is  surrounded  by  four  lobes.  In  the 
Ptychopteridae  the  very  reduced  disk  is  borne  at  the  tip  of  a  slender, 
retractile  breathing  tube.  In  the  Tanyderidae  the  condition  is  somewhat 
similar,  but  here  the  disk  is  larger  and  is  surrounded  by  five  lobes  at  the 
tip  of  a  long,  stout,  non-retractile  breathing  tube. 

The  spiracles  vary  greatly  in  size  from  very  large  to  small  and  vestigial, 
or  they  may  even  be  lacking  in  some  species  of  Antocha.  They  consist 
of  an  apparently  uniform  middle  piece  surrounded  by  a  radially  folded 
margin,  or  ring,  of  various  widths,  called  the  stigmal  ring.  Many  authors 
(De  Meijere,  Mik,  Mtiggenberg,  Brown,  Keilin,  and  others)  hold  that 
the  middle  piece  is  an  imperforate  chitinized  plate  and  that  respiration 
takes  place  thru  the  stigmal  ring.  Gerbig  (1913),  however,  shows  that 
the  middle  piece  is  split  across  the  disk,  .the  cleft  being  closed  by  two 
overlapping  membranes.  Directly  behind  the  spiracles  the  tracheae 
enlarge  into  the  felt  chamber,  whose  walls  are  provided  with  long,  branched,' 
treelike  structures,  the  branches  apparently  anastomosing.  Surrounding 
the  felt  chamber  in  many  larvae  are  dense  masses  of  air  tubes,  which 
make  up  the  tracheal  lungs.  These  tubes  are  arranged  in  bundles,  which 
arise  in  special  cavities  of  the  felt  chamber;  thus,  in  Tipula  paludosa, 
there  are  about  fifty  bundles,  each  of  about  twenty  tubules,  making 
a  total  of  one  thousand  of  these  air  canals  (Gerbig). 

The  early  stages  of  the  larva  are  quite  different  from  the  later  develop- 
mental stages,  as  Gerbig  (1913:137-140),  working  on  Tipula  paludosa, 
has  well  shown.  The  prominent  six-lobed  spiracular  disk  of  the  more 
matured  larva  is  represented  in  the  first  developmental  stage  by  four 
heavily  chitinized  projections,  which  bear  but  few  bristles  on  their  outer 
margin.  The  dorsal  lobes  are  not  evident,  but  are  replaced  by  eight 
branched  bristles,  about  equidistant  from  one  another.  The  spiracles 
are  oval,  not  circular  as  in  the  grown  larva,  and  project  a  little  beyond 
the  level  of  the  disk.  The  writer  has  noted  several  first-stage  larvae 
with  an  appearance  almost  as  described  but  showing  several  points  of 
difference.  The  immature  larvae  of  Phalacrocera  are  described  elsewhere 
in  this  work  (page  963). 


748  CHARLES  PAUL  ALEXANDER 

Anal  gills.— In  the  Ptychopteridae  and  in  the  Tanyderidae  there  is 
but  a  single  pair  of  anal  tracheal  gills.  In  the  former  these  are  very  small 
and  are  cylindrical,  in  the  latter  they  are  large,  branched,  fanlike  struc- 
tures. In  the  Tipulidae  the  anal  gills  number  from  four  in  most  Lim- 
nobiinae  to  six  or  eight  in  the  Tipulinae.  The  increase  in  number  is  brought 
about  by  a  division  of  the  original  gill  of  each  side.  In  generalized  forms, 
as  Antocha,  Pedicia,  and  others,  the  anal  gills  are  constricted  into  segments, 
the  apical  ones  being  more  or  less  telescopic  into  the  preceding  ones.  As 
a  rule  the  anal  gills  are  entirely  or  almost  entirely  retractile  within  the 
body. 

In  the  Tipulinae  the  gills  vary  with  the  genera,  being  blunt  and  con- 
structed for  propulsion  in  the  wood-inhabiting  species,  such  as  those  of 
Ctenophora,  Tanyptera,  Tipula,  and  other  genera.  In  the  semi-aquatic 
species  of  Tipula  the  number  of  gills  varies  from  four  to  eight.  In  the 
latter  case  each  of  the  four  principal  gills  is  deeply  bifid  and  the  gills  are 
arranged  transversely,  as  in  T.  ignobilis]  in  species  with  six  gills  the  posterior 
branches  of  the  posterior  gills  are  atrophied  as  a  rule;  in  other  species, 
which  have  but  four  gills,  the  four  anterior  branches  are  preserved,  the 
posterior  pair  being  usually  atrophied.  In  Longurio  and  Aeshnasoma 
the  four  anal  gills  are  pinnately  branched,  each  with  about  six  lateral 
branches.  A  similar  condition  is  found  in  the  Tanyderidae. 

The  head 

The  primitive  crane-fly  head  was  undoubtedly  of  the  eucephalous, 
non-retractile  type,  as  found  in  the  Ptychopteridae,  the  Rhyphidae,  and 
other  families.  The  retracted  head  capsule  of  the  Tipulidae  is  a  derived 
condition. 

The  massive,  compact  capsule  is  found  in  all  of  the  lower  groups  of  the 
three  subfamilies  of  the  Tipulidae.  Such  a  head  is  easily  derivable  from 
the  condition  in  the  Rhyphidae  or  in  the  Tanyderidae,  for  instance.  The 
dorsal  median  sclerite,  the  prefrons,  is  almost  as  large  and  conspicuous 
in  Ula  as  in  the  eucephalous  families.  The  lateral  plates  that  constitute 
the  capsule  are  shaped  like  a  mussel  in  the  generalized  groups,  with  the 
posterior  incisions  shallow.  In  the  more  specialized  forms,  with  the 
capsule  greatly  dissected,  the  two  plates  of  either  side  are  entirely  sepa- 
rated—  the  innermost,  next  to  the  prefrons,  being  the  internal-lateral 
of  De  Meijere,  and  the  outermost  being  his  external-lateral.  The  prefrons 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  749 

is  found  in  all  forms  with  the  capsule  compact  and  massive,  and  is  even 
preserved  in  many  species  with  the  dissected  capsule.  In  many  cases, 
however,  it  is  firmly  fused  with  the  internal-lateral  plates;  or,  in  some 
cases,  all  the  plates  of  the  capsule  are  firmly  united  and  their  individual 
limits  are  ill-defined.  In  the  generalized  forms  the  clypeus  and  the 
labrum  are  entirely  distinct  from  each  other,  while  in  other  forms  the  two 
lose  their  individuality  by  fusion. 

The  sclerites  of  the  generalized  primitive  head  bear  bristles,  or  setae, 
of  various  types  and  arrangement.  In  the  Tanyderidae  the  punctures 
are  multisetose;  in  most  Ptychopteridae  the  setae  are  branched  or  some- 
what plumose. 

The  literature  on  the  head  capsule  and  the  mouth  parts  is  as  follows: 

Ptychoptera.     De  Meijere,  1916:188-191,  figs.  14-20. 
Bittacomorpha.     Hart,  1898  [1895]:  192,  pi.  5. 
Trichocera.     De  Meijere,  1916:191-194,  figs.  21-23. 
Limnobia.     De  Meijere,  1916:198-201. 
Dicranomyia.     De  Meijere,  1916: 197-198,  figs.  32-35. 
Dicranoptycha.     Alexander,  1919 b: 71,  figs.  2,  3. 
Elliptera.     Mik,  1886 b: 339,  pi.  6. 
Ula.     Alexander,  1915  a :  5-6,  pi.  1. 

Dactylolabis.     Mik,  1894:261-266,  pi.  2;  Nowicki,  1867:341  (as  Rhicnoptila). 
Limnophila.     De  Meijere,  1916:204-206,  figs.  49-51. 
Eriocera.     Alexander  and  Lloyd,  1914:21-22,  pi.  1. 
Hexatoma.     Alexander,  1915  c:  146,  pi.  1. 
Penthoptera.     Alexander,  1915 c:  155,  pi.  1. 
Tricyphona.     De  Meijere,  1916: 195-196,  figs.  29-31. 
Pedicia.     Needham,  1903:286,  fig.  19. 
Dicranota.     Miall,  1893:  237-238,  pi.  10. 
Ormosia.     De  Meijere,  1916:201-204,  figs.  37-47. 
Gnophomyia.     Malloch,  19 15-17 b: 230-231,  pi.  34,  fig.  10. 
Chionea.     Brauer,  Egger,  and  Frauenfeld,  1854:614,  pi.  11. 
Phalacrocera.     Miall  and  Shelf ord,  1897:344-345;  Bengtsson,  1897. 
Cylindrotoma.     Cameron,  1918. 

Liogma.     Miiggenberg,  1901;  Alexander,  1914:  111,  pi.  1. 
Ctenophora.     Anthon,  1908:544. 
Prionocera.     Malloch,  19 15-17 b:  199,  pi.  32,  figs.  1-3. 
Holorusia.     Comstock  and  Kellogg,  1904:55,  61-62. 
Tipula.     Vimmer,  1906,  and  1911:1-6. 

In  addition  to  the  preceding,  the  genera  Helobia  (Beling,  Hart,  Malloch),  Rhaphidolabis 
(Needham),  and  Tanyptera  have  been  discussed  less  fully. 

All  of  the  above-named  genera  are  considered  in  this  paper,  and  the 
head  capsule  and  mouth  parts  of  the  following  genera  are  described 
also:  Protoplasa  (supposition),  Bittacomorphella,  Rhipidia.  Rhamphidia, 
Antocha,  Epiphragma,  Pseudolimnophila,  Pilaria,  Ulomorpha,  Elephant- 
omyia,  Teucholabis,  Gonomyia,  Erioptera,  Molophilus,  Adelphomyia 


750  CHARLES  PAUL  ALEXANDER 

(supposition),  Rhaphidolabina,  Oropeza,  Longurio,  Aeshnasoma.  The 
descriptions  for  the  genera  are  either  entirely  new  or  else  the  characters 
have  heretofore  been  insufficiently  described  and  figured.  The  details 
of  structure  of  the  head  capsules  of  nearly  fifty  genera  of  crane-flies  are 
now  known. 

The  labrum  and  the  epipharynx. —  The  labrum  is  preserved  in  all  crane- 
fly  larvae.  It  is  usually  broadly  transverse  or  oval.  The  dorsal  surface 
is  clothed  with  short  hairs,  these  being  longer  on  the  anterior  margin, 
which  is  often  provided  also  with  a  limited  number  of  sensory  bristles, 
or  papillae.  The  lateral  regions  on  the  ventral,  or  epipharyngeal,  side 
often  bear  long  tufts  of  hairs.  The  epipharynx  proper  is  variously  armed 
in  the  different  groups  of  crane-flies. 

The  labium. — The  labial  region  of  the  capsule  is  of  vast  importance  in 
classification,  and  the  location  of  the  constituent  parts  should  be  early 
appreciated.  The  confusion  in  terminology  of  the  dipterous  larval  labium 
has  been  partly  outlined  by  De  Meijere  (1916:253).  The  principal 
synonyms  are  as  follows: 

Submentum. 

Mentum  —  The  pharyngeal  plate,  or  lower  lip  (Meinert) ;  submentum  (Miall) ;  ectolabium 
(Bengtsson);  labial  plate. 

Prementum  —  Mentum  (Miall) ;  ectolabium  (Keilin) ;  endolabium  (Holmgren,  Vimmer, 
Bengtsson). 

Hypopharynx  —  Labium  (Meinert);  mentum  (Miall,  in  Dicranota);  endolabium  (Keilin). 

The  submentum  is  represented  by  a  narrow  transverse  strip  in  the 
eucephalous  groups  of  crane-flies,  being  well  shown  in  Ptychoptera.  The 
mentum  is  the  usually  chitinized  anterior-ventral  plate  of  the  capsule. 
In  Ptychoptera  it  is  margined  anteriorly  with  about  twenty  comblike 
teeth.  In  Bittacomorpha  it  is  more  or  less  distinctly  bilobed,  but  is 
untoothed.  In  the  generalized  Tipulidae  it  is  indistinctly  divided  into 
an  outer  plate  and,  immediately  dorsad  of  this  and  closely  applied  to  or 
fused  with  it,  an  inner  plate.  The  outer  plate  furnishes  the  apical  median 
tooth  of  the  mentum  and  in  some  cases  an  additional  tooth  on  either 
side.  The  inner  plate  adds  a  varying  number  of  teeth  to  the  mental 
plate,  from  two  in  Epiphragma  to  as  many  as  ten  or  twelve  in  some  species 
of  Limnobia.  The  chitinized  plate  is  completely  divided  medially  in 
some  Eriopterini  (Molophilus  and  some  Erioptera),  in  the  Pediciini,  and 
in  some  Hexatomini  (Pseudolimnophila).  In  this  case,  each  half  is 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  751 

directly  continuous  with  the  ventral  bars  of  the  head  capsule.  In  the 
Limnophilaria  the  mentum  is  represented  by  a  narrow,  transverse, 
chitinized  bar,  which  is  delicately  grooved  with  parallel  lines.  The 
mentum  is  lacking  in  chitinized  parts,  or  apparently  so,  in  many  Eriop- 
terini  and  in  the  higher  Hexatomini. 

Directly  behind  the  mentum  is  the  prementum,  rarely  prominent  and 
bearing  the  two  small  palpi  when  they  are  preserved,  as  in  the  Ptychopter- 
idae.  The  hypopharynx  lies  immediately  dorsad  of  the  prementum.  In 
the  Ptychopteridae  this  is  an  enlarged,  two-horned,  fleshy  lobe  which 
is  difficult  to  study.  In  the  Tipulidae  the  generalized  condition,  such 
as  occurs  in  the  Limnobiini  and  the  lower  Hexatomini,  consists  of  two 
collar-like  chitinized  plates  whose  anterior  margins  bear  from  ten  to 
fifteen  or  more  teeth.  In  the  Tipulini  this  plate  is  narrow,  with  the 
anterior  margin  few-toothed,  there  usually  being  five  or  in  some  cases 
six  teeth.  In  the  Pediciini  the  hypopharynx  is  labriform;  in  many 
Eriopterini  it  is  cushion-shaped  and  covered  with  numerous  blunt  setae. 

The  antennae. —  The  antennae  are  borne  on  the  anterior  lateral  margin 
of  the  capsule,  laterad  of  the  labrum.  They  are  one-  or  two-segmented, 
and  in  most  cases  have  one  or  more  apical  papillae.  De  Meijere  and 
others  hold  that  the  principal  apical  papilla  constitutes  an  additional 
antennal  segment.  In  some  cases,  however,  as  in  Ula  for  example,  two 
or  even  more  papillae  are  found  which  are  very  similar  to  one  another 
except  for  slight  degrees  of  size.  It  is  difficult  in  such  cases  to  hold  any 
one  of  these  papillae  as  a  true  segment.  The  writer  prefers  to  consider 
them  as  apical  sensory  papillae,  and  they  are  so  treated  in  this  paper. 

The  primitive  antennae  are  not  greatly  elongated,  and  are  usually 
short-cylindrical  or  subglobular  (Ula).  An  auditory  plate,  circular  in 
outline  and  lying  on  the  face  of  the  basal  segment,  is  almost  always  present. 
The  apical  papilla  in  the  Limnophilaria  and  the  Eriopterini  is  egg-shaped, 
or  oval,  and  is  delicately  sculptured  by  a  network  of  apparently  impressed 
lines.  In  many  species,  as  for  example  those  of  the  higher  Hexatomini, 
the  apical  papillae  are  very  long,  tapering  to  the  tips.  In  many  Limno- 
biini, as  for  example  Limnobia,  and  in  many  species  of  Tipula  and  related 
forms,  the  apical  papilla  is  button-shaped,  or  disklike,  and  is  often  very 
reduced. 

The  mandible. —  In  the  primitive  type,  such  as  occurs  in  the  eucephalous 
groups  and  in  the  generalized  Tipulidae,  the  mandible  is  rather  com- 


752  CHARLES  PAUL  ALEXANDER 


plicated,  usually  having  a  considerable  number  of  teeth  on  the  cutting 
edge  and  a  distinct  sub-basal  arm  (prostheca)  or  prosthecal  tuft  of  hairs. 
At  the  same  time  there  are  usually  one  or  more  setae  on  the  outside  of 
the  mandible,  near  the  "  heel,"  or  base,  of  the  scrobal  region.  In  the 
generalized  tipulid  type  there  are  two  apical  teeth  and  two  rows  of  cutting 
teeth.  In  other  cases,  as  in  the  Limnobiini,  there  are  but  a  single  dorsal 
tooth,  a  single  apical  tooth,  and  a  varying  number  of  cutting  teeth  on 
the  ventral  face;  in  some  species  of  Limnobia  the  number  of  teeth  in  the 
ventral  cutting  row  is  six  or  seven.  In  the  Tipulini  the  mandible  is  often 
reduced  in  size  (as  in  Tanyptera),  there  being  but  a  single  dorsal  and 
a  single  ventral  tooth  in  addition  to  the  apical  point;  the  prostheca, 
however,  is  usually  well  developed.  In  the  Eriopterini  but  one  row  of 
teeth,  the  ventral,  is  strongly  developed.  In  the  Pediciini  and  the  higher 
Hexatomini,  the  mandible  is  elongate  and  sickle-shaped,  with  the  few 
teeth  on  the  cutting  edge  at  about  midlength.  In  Ulomorpha  the  mandible 
is  hinged  at  about  midlength,  the  basal  part  being  hollowed  out  on  the 
inner  face  to  receive  the  blade  in  a  position  of  rest. 

The  maxillae. —  The  maxillae  are  usually  of  simple  form.  In  the 
generalized  types,  such  as  the  eucephalous  families  and  the  lower  groups 
of  the  subfamilies  of  Tipulidae,  they  consist  of  a  large  and  distinct  trans- 
versely triangular  cardo,  a  conspicuous  stipes,  and  distinct  inner  ar,d 
outer  apical  lobes.  In  the  Pediciini  the  lobes  are  separate  in  the  supposed 
Adelphomyia  larva  but  are  fused  together  in  the  highest  types  (Dicranota) . 
In  the  Hexatomini  the  lobes  are  reduced  to  a  single  long,  flattened  blade. 
The  outer  and  inner  lobes  are  usually  densely  hairy,  especially  at  the  tip 
and  on  the  inner  face  of  the  latter.  The  palpus  is  uni-articulate  and 
antenniform  in  the  generalized  forms,  is  short-cylindrical  with  a  varying 
number  of  tiny  sense  pegs  at  the  apex,  and  bears  on  its  face  a  circular 
auditory  plate  similar  to  that  of  the  antenna.  The  armature  of  the  inner 
lobes  is  varied,  in  some  species  being  provided  with  stiff,  comblike  setae, 
or  projections. 

The  pupa 
General  features 

The  pupa  of  the  crane-fly  is  of  the  so-called  "free,"  or  "  mummy," 
type.  In  only  a  few  cases  does  the  larval  skin  adhere  to  the  end  of  the 
abdomen  (Dactylolabis,  Cylindrotoma,  and  some  other  forms.)  The  head 
is  usually  small,  and  is  ventral  in  position.  Immediately  behind  the 


THE  CKANE-FLIES  OF  NEW  YORK  —  PART  II  753 

head  on  the  pronotum  are  the  two  breathing  horns,  which  are  variously 
developed  in  the  different  tribes  and  genera  and  furnish  invaluable  bases 
for  classification.  The  leg  sheaths  usually  far  exceed  the  short  or  but 
moderately  elongate  wing  sheaths.  The  abdominal  segments  are  often 
provided  with  a  subterminal  armature  of  stiff  setae,  or  spines.  At  the 
posterior  end  of  the  body,  the 'last  two  segments  (cauda)  are  variously 
modified  to  inclose  the  sexual  organs  of  the  adult  flies. 

The  head 

The  head  is  usually  small  and  flattened,  occupying  the  anterior  ventral 
part  of  the  body.  The  eyes  differ  in  size  in  the  various  groups,  in  some 
(Erioptera,  Elephantomyia)  being  larger  in  the  male  than  in  the 
female ;  in  the  male  sex  they  are  approximated  on  the  median  line  above 
or  beneath. 

The  front  between  the  eyes  is  usually  narrowed  behind,  delimited  by 
the  inner  margin  of  the  compound  eye,  narrowed  at  the  posterior  end, 
and  bluntly  rounded  or  pointed  at  the  apex.  This  part  is  described 
herein  as  the  Idbrum,  or  labral  sheath.  It  contains  the  fronto-clypeus 
and  the  labrum  of  the  imago.  At  its  tip  it  bears  two  more  or  less  diver- 
gent lobes,  these  being  in  some  cases  closely  approximated  so  as  to  appear 
as  a  single  lobe;  these  are  herein  termed  the  labial  lobes  or  sheaths,  and  they 
contain  the  so-called  paraglossae  of  the  adult  fly. 

On  either  side  of  the  labral  sheath,  and  usually  divergent  and  lying 
along  the  posterior  margin  of  its  face,  are  the  sheaths  of  the  maxillary 
palpi,  which  in  most  cases  extend  beyond  the  knee  joint  of  the  fore  legs. 
In  almost  all  the  Limnobiinae  these  are  short  and  stout  and  almost 
straight,  but  in  the  majority  of  the  tipuline  forms  they  are  curved  at 
their  tips,  which  in  most  species  are  actually  recurved  and  offer  an  easy 
means  of  distinguishing  members  of  this  subfamily.  In  many  of  the 
Limnobiini  the  margins  of  the  cheeks  project  as  flattened  ledges  overlying 
the  joint  of  the  fore  legs. 

The  antennae  arise  from  above  or  between  the  eyes  and  bend  laterad 
and  thence  caudad  around  the  eyes,  in  some  forms,  such  as  Elephant- 
omyia and  the  males  of  other  species,  lying  across  the  face  of  the  eyes. 
The  antennae  usually  end  just  beyond  the  roots  of  the  wings,  but  in  the 
males  of  some  species  (Eriocera)  they  extend  beyond  the  tips  of  the  wings. 
The  basal  segments  (scape)  of  the  antennal  sheath  are  often  armed  with 

3 


754  CHARLES  PAUL  ALEXANDER 

spines  or  tubercles,  and  very  often  the  organ  is  angulated  at  the  end  of 
each  segment  of  the  adult  antenna  inside. 

At  its  vertex,  between  or  just  dorsad  of  the  antennal  bases,  the  head 
may  bear  a  crest  which  is  usually  bilobed  and  setiferous.  In  some  species 
this  cephalic  crest  is  quadrituberculate,  there  being  a  smaller  secondary 
crest  behind  or  before  the  primary  one.  -In  the  Tipulini  the  crest  is  very 
inconspicuous  and  but  weakly  setiferous.  In  most  of  the  Limnobiini 
it  is  lacking  or  nearly  so. 

The  head  may  be  variously  armed  with  spines,  tubercles,  or  setae;  in 
Eriocera  spinosa,  for  example,  there  are  spines  or  strong  tubercles  on  the 
antennal  scape,  on  the  clypeal  region,  and  even  on  the  face  of  the  eye. 
In  some  cases  there  are  setae  on  the  front  between  the  eyes,  on  the 
clypeus,  and  on  the  cheek. 

The  thorax 

The  pronotum  of  the  thorax  is  small.  The  ventral  part  is  closely 
applied  to  the  head  and  often  has  small  setiferous  tubercles  close  to  the 
breathing  horns.  The  pronotal  breathing  horns  are  variously  developed 
in  the  different  tribes  and  genera,  and  are  discussed  here  in  general  terms 
only. 

Many  species  are  propneustic,  the  pronotal  horns  alone  being  functional. 
Other  species  (in  Hexatomini  and  Eriopterini)  are  peripneustic,  the  second 
to  the  seventh  abdominal  segments  being  provided  with  functional  lateral 
spiracles  in  addition  to  the  breathing  horns;  other  pupae  have  lateral 
abdominal  spiracles,  but  in  most  cases  they  are  merely  vestigial.  Some 
pupae  are  amphipneustic,  there  being  in  addition  to  the  breathing  horns 
a  conspicuous  pair  of  spiracles  on  the  dorsum  of  the  eighth  abdominal 
segment  (Rhamphidia,  Ula,  Epiphragma;  in  the  typical  species  of  Lim- 
nobia  these  are  present  but  they  are  small  and  are  probably  nonfunctional) . 

In  the  Ptychopteridae  the  breathing  horns  are  very  unequally  developed, 
one  being  enormously  elongated  and  filiform  while  the  other  is  abortive. 
In  some  Tipulini  (Longurio,  Prionocera,  Tipulodina)  the  horns  are  like- 
wise greatly  elongated,  but  in  these  cases  they  are  shorter  than  the 
body  and  are  approximately  subequal  in  size,  or  at  least  are  not  so 
disproportionately  unequal. 

In  the  Limnobiini  the  breathing  horns  are  usually  stout  and  broad, 
in  the  typical  Limnobaria  (Limnobia,  Dicranomyia)  being  subquadrate, 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  755 

subcircular,  or  even  broader  than  long.  In  Elliptera  they  are  earlike  and 
very  large.  In  Antocha  a  unique  condition  is  found,  in  that  the  apparatus 
is  divided  into  eight  stout  filaments.  In  the  Eriopterini  and  the  Hexatomini 
the  horns  are  usually  long  and  cylindrical  or  slightly  flattened,  and  straight 
or  but  slightly  curved.  There  are  numerous  deviations  from  type,  how- 
ever. In  Gnophomyia  and  some  species  of  Gonomyia  (alexanderi  and 
kansensis,  for  example),  the  horns  are  trumpet-shaped;  in  Gonomyia  sul- 
phur ella  they  are  compressed  and  flattened  into  fanlike  structures.  In 
several  widely  separated  paludicolous  genera,  such  as  Ulomorpha,  Pseu- 
dolimnophila,  and  Prionocera,  the  breathing  horns  are  split  at  their  tips 
into  two  conspicuous  flaps  with  fimbriate  margins,  an  obvious  adaptation 
to  an  existence  in  mud.  Often  the  breathing  horns  are  very  small  (as  in 
Elephantomyia,  Teucholabis,  and  Trichocera)  or  even  microscopic 
(Dicranoptycha).  Limnophila  hyalipennis  (Zett.)  is  described  by  Beling 
(1886 : 198-199)  as  lacking  the  breathing  horns,  but  this  is  presumably  an 
error  of  observation  which  may  be  due  to  a  defect  in  the  material  studied, 
or  possibly  the  horns  are  sessile  or  reduced  as  in  Dicranoptycha.  In 
some  species  (Erioptera,  Eriocera)  the  horns  are  stout  at  the  base  but 
taper  rapidly  to  the  acute  points.  In  many  Pediciini  the  tips  of  the 
horns  are  expanded  and  usually  obliquely  truncated,  with  a  row  of 
breathing  pores  around  the  margin.  The  tipuline  breathing  horns  are 
remarkably  uniform  in  general  structure,  being  usually  elongated  cylindrical 
in  shape,  of  moderate  length,  and  with  the  tips  slightly  expanded.  The 
most  conspicuous  deviations  from  type  are  the  short,  flattened,  and  some- 
what clavate  horns  of  Tanyptera. 

The  thoracic  mesonotum  is  very  convex,  or  gibbous  in  many  species 
(Limnobiini  and  some  Hexatomini).  In  other  species  it  is  declivitous, 
with  a  high  transverse  crest  (Eriopterini).  This  crest  may  be  armed 
with  numerous  small  spines  (Helobia);  from  two  to  four  powerful  hooks 
(Molophilus,  some  species  of  Ormosia,  some  species  of  Erioptera,  Teucho- 
labis); from  six  to  eight  tubercles  set  with  stiff  bristles  (Gonomyia);  or 
abundant  small  setiferous  tubercles  on  either  side  of  the  median  line, 
these  being  less  conspicuous  along  the  shoulder  (most  species  of 
Ormosia  and  of  Erioptera).  In  Eriocera  longicornis  there  is  a  median 
spine  or  tubercle  on  the  scutellum.  In  some  Cylindrotominae  the 
metanotum  bears  spines.  The  extreme  lateral  or  ventral  margin  is 
produced  into  a  blunt  or  somewhat  pointed  angle  just  above  the  wing 


756  CHARLES  PAUL  ALEXANDER 

root;  this  is  usually  armed  with  from  one  to  three  setae.  Various  other 
setae  usually  occur  on  the  mesonotum. 

The  leg  sheaths  lie  along  the  ventral  side  of  the  pupa.  There  is,  in  the 
various  groups,  a  considerable  difference  in  the  length  of  the  sheaths, 
their  position,  and  the  relative  arrangement  of  the  ends  of  the  tarsal 
segments.  In  the  Bittacomorphinae,  the  fore  legs  overlie  the  middle 
legs.  In  the  Rhyphidae  the  fore  legs  overlie  the  middle  pair,  which, 
in  turn,  overlie  the  hind  legs.  In  the  Ptychopterinae  and  in  the  Tipulidae, 
all  three  pairs  of  sheaths  lie  parallel  along  the  midventral  area,  those 
of  the  hind  legs  being  outermost  along  the  wing  sheaths,  and  those  of 
the  fore  legs  being  on  the  inside  and  contiguous.  In  Gnophomyia  the 
leg  sheaths  are  very  short,  extending  but  little  beyond  the  wing  tips  and 
reaching  only  the  end  of  the  second  abdominal  segment.  In  other  crane- 
flies  they  are  longer,  in  some  species  reaching  the  end  of  the  fifth  abdominal 
segment.  Very  often  the  tips  of  the  tarsi  are  on  a  level.  In  the  Limno- 
biini  the  hind  tarsi  are  usually  a  little  shorter,  and  the  fore  tarsi  are  a  little 
longer,  so  that  the  ends  of  the  sheaths  form  a  blunt  V.  In  the  Eriopterini 
the  middle  legs  are  usually  (except  in  Gnophomyia)  much  shorter  than 
the  other  legs;  in  Helobia,  moreover,  the  hind  legs  are  conspicuously 
longer  than  the  fore  legs,  so  that  all  three  sets  end  at  different  levels. 
A  somewhat  similar  and  parallel  development  is  found  in  the  tipuline 
forms. 

The  wing  sheaths  are  moderately  broad,  usually  ending  opposite  or 
a  little  beyond  the  apex  of  the  second  abdominal  segment.  The  venation 
of  the  various  species  often  shows  clearly  and  with  little  distortion  on 
the  wing  pad,  and  in  such  cases  it  is  of  very  great  help  in  locating  certain 
genera.  Species  with  a  heavy  pattern  in  the  adult  wings,  such  as  those 
of  the  genera  Discobola  and  Epiphragma,  show  this  pattern  on  the  wing 
pad  in  the  older  pupae,  and  this  helps  to  confirm  their  identity. 

The  abdomen 

The  abdomen  consists  of  apparently  nine  segments,  the  last  two  being 
closely  approximated  or  fused  to  form  the  cauda.  The  segments  are 
usually  plainly  divided  into  a  broad  dorsal  (tergal)  and  ventral  (sternal) 
region,  and  a  much  narrower  and  more  restricted  lateral  (pleural)  area. 
The  segments  beyond  the  first  are  variously  subdivided  into  rings,  or 
annuli,  by  false  constrictions,  there  being  usually  one,  but  in  some  cases 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  757 

two  or  more,  basal  rings  and  a  much  broader  posterior  ring.  The  segments 
are  variously  armed  with  setae,  or  spines.  In  the  Limnobiini  and  some 
Hexatomini  these  occur  on  the  basal  annulus,  and  consist  of  a  transverse 
welt  which  is  margined  (as  in  Antocha  and  Elliptera)  or  covered  with 
abundant  microscopic  chitinized  points.  In  some  Pediciini  the  setae 
occupy  both  the  basal  and  the  posterior  rings  of  the  intermediate  abdominal 
segments.  In  the  Eriopterini  and  most  Hexatomini,  as  well  as  in  the 
tipuline  forms,  the  setae  occur  on  the  posterior  ring,  just  before  the  caudal 
margin,  and  consist  of  a  single  transverse  row  of  small  spines.  In  the 
Cylindrotominae  the  segments  are  often  armed  with  prominent  elongate 
body  projections  (Liogma,  Triogma).  In  Phalacrocera  elongate  lobes 
are  confined  to  the  posterior  segments  of  the  -abdomen.  In  the  Pty- 
chopteridae  the  segments  are  densely  covered  with  transverse  rows  of 
long  tubercles  which  are  replaced  by  chitinized  plates  near  the  posterior 
margin  of  the  segments.  In  Bittacomorpha  these  tubercles  are  stellate 
at  their  tips. 

The  dorsum  of  the  cauda  in  most  crane-flies  is  armed  with  four,  five, 
or  six  conspicuous  lobes  which  are  often  spinous  at  or  near  their  tips. 
These  represent  the  lobes  that  surround  the  spiracular  disk  of  the  larva, 
and  their  number  generally  corresponds  to  the  larval  condition.  Thus, 
in  the  Ptychopteridae  there  is  a  single  lobe,  in  the  Pediciini  there  are  two, 
and  in  the  Hexatomini  and  the  Eriopterini  there  are  four  or  five.  In  the 
Tipulinae  there  are  often  but  four  evident  lobes,  the  dorsal  pair 
of  the  larva  not  being  represented.  In  some  generalized  groups,  as 
the  Limnobaria,  the  Rhamphidaria,  and  others  as  indicated  else- 
where in  this  paper,  the  larval  spiracles  persist  in  a  more  or  less  functional 
condition. 

Abdominal  spiracles. —  In  the  Eriopterini  and  the  Hexatomini  there  is 
a  distinct  and  apparently  functional  spiracle  on  each  pleurite  of  the  second 
to  the  seventh  abdominal  segments,  located  near  the  base  of  the  posterior 
ring.  In  the  Eriopterini  these  spiracles  are  small  and,  in  the  cast  pupal 
skin,  appear  tubular  or  elongate.  Smaller  vestigial  spiracles  are  evident 
in  most  genera  of  the  Tipulidae. 

As  has  been  mentioned  earlier  in  this  paper,  in  certain  primitive  genera 
of  Tipulidae  —  such  as  Rhamphidia,  Ula,  Epiphragma,  and  to  a  lesser 
extent  those  of  Limnobaria  —  there  is  a  pair  of  rather  large  spiracles 
on  the  dorsum  of  the  eighth  segment.  In  Dactylolabis  cubitalis  the  lateral 


758  CHARLES  PAUL  ALEXANDER 

abdominal   spiracles  are   protuberant,   notably  the  pair  on   the   second 
segment. 

The  genitalia. —  The  genital  sheaths  conceal  the  adult  organs  within. 
In  the  male  the  valves  are  usually  subequal  or  the  sternal  valves  are 
a  little  the  longer.  In  the  female  the  sheaths,  or  acidothecae,  of  the 
ovipositor  usually  have  the  dorsal  valves  considerably  longer  than  the 
sternal  valves  and  distinctly  upturned  at  their  tips.  In  the  few  groups 
with  fleshy  ovipositors,  the  female  cauda  is  very  difficult  to  distinguish 
from  the  cauda  of  the  male.  The  valves  are  variously  armed  with  spines 
or  lobes,  at  or  before  the  tips. 

INTERNAL   MORPHOLOGY 

The  internal  morphology  and  anatomy  of  the  crane-fly  larva  are  here 
considered  in  general  terms  only.  The  two  genera  that  have  been  studied 
in  greatest  detail  by  the  writer  are  Eriocera  and  Antocha.  The  most 
important  literature  on  the  various  systems  of  the  body  is  cited  in  order 
to  make  reference  to  it  more  easily  available. 

The  digestive  system 
The  most  important  literature  on  the  digestive  system  is  as  follows : 

Ptychoptera.     Grobben,  1876;  Van'Gehuchten,  1890. 

Chionea.     Brauer,  Egger,  and  Frauenfeld,  1854:613-614,  pi.  11,  fig.  4. 

Dicranota.     MiaU,  1893:242-245. 

Phalacrocera.     Miall  and  Shelford,  1897:347-351;  Bengtsson,  1897. 

Ctenophora.    Anthon,  1908:546-551,  figs.  12-35. 

Holorusia.     Kellogg,  1901  a;  Comstock  and  Kellogg,  1904:56-57. 

Herbivorous  larvae  (Tipula,  Holorusia,  Ctenophora,  Phalacrocera) 

In  the  larva  of  herbivorous  species  the  alimentary  canal  extends  the 
length  of  the  body  as  an  almost  straight  tube  inclosed  by  the  coiled, 
perforated  sheets  of  adipose  tissue.  The  esophagus  is  slender,  opening 
into  the  hypopharynx.  The  pro  ventriculus  has  a  large  esophageal  in  vagi- 
nation  at  its  anterior  end.  The  ventriculus  has  at  its  anterior  end  four 
elongated  ventricular  caeca,  these  being  of  two  distinct  lengths  in  pairs 
(didynamous)  in  Ctenophora,  and  all  four  alike  in  Holorusia.  In  Cteno- 
phora there  are  four  small,  pocket-like  gastric  caeca  near  the  posterior  end 
of  the  ventriculus,  which  are  not  mentioned  as  occurring  in  Holorusia. 
At  the  juncture  of  the  ventriculus  and  the  small  intestine  are  the  four 


THE  CBANE-FLIES  OF  NEW  YORK  —  PART  II  759 

coiled  malpighian  tubules.  Behind  the  ventriculus  lies  the  small  intestine, 
with  a  caliber  much  smaller  than  that  of  the  ventriculus.  At  the  union 
of  the  large  and  the  small  intestine  is  a  prominent  diverticulum  of  very 
large  size  (in  Ctenophora  three  times  as  large  as  in  Holorusia).  The 
large  intestine  gradually  dilates  behind  to  form  the  rectum. 

Phalacrocera  has  the  ventriculus  without  caeca  and  the  hind  intestine 
without  a  diverticulum,  altho  it. is  decidedly  a  herbivorous  genus.  The 
condition  in  Chionea  is  almost  the  same.  In  Phalacrocera,  and  probably 
in  most  other  larvae,  a  peritrophic  membrane  (a  secretion  of  the  epithelium 
which  incases  the  food  and  keeps  it  from  direct  contact  with  the  epithe- 
lium) is  present. 

There  are  four  malpighian  tubules  in  all  the  Tipulidae  that  have  been 
thoroly  studied  (Chionea,  Eriocera,  Dicranota,  Phalacrocera,  Ctenophora, 
Holorusia,  Tipula,  and  a  few  others).  In  the  Rhyphidae,  likewise,  there 
are  four  tubules  in  both  the  larval  and  adult  stages.  In  the  Ptychop- 
teridae,  however,  there  are  five  tubules,  as  in  the  Culicidae,  the  Psy- 
chodidae,  and  related  families.  This  might  indicate  some  basis  for  placing 
the  Ptychopteridae  in  close  proximity  to  the  Culicidae,  as  has  been  done 
by  Lameere  (1906). 

The  salivary  glands  consist  of  two  large  coiled  tubules  which  pass  into 
a  collecting  duct  and  unite  to  form  the  common  collecting  duct  that  opens 
at  the  base  of  the  hypopharynx. 

Carnivorous  larvae  (Dicranota,  Eriocera) 

The  alimentary  canal  in  the  carnivorous  type  of  crane-flies  is  a  short, 
straight  tube,  with  a  minimum  of  appendages  such  as  caeca  and  diver- 
ticula.  In  Eriocera  the  esophagus  is  very  long,  and  the  ventriculus  is 
short  and  without  caeca.  The  malpighian  tubules  are  yellow  at  their 
origin,  soon  passing  into  an  orange-brown  color.  The  intestine  is  short 
and  straight,  without  a  diverticulum. 

The  salivary  glands  in  the  newly  killed  larva  of  Eriocera  spinosa  are 
very  conspicuous  by  their  opalescent  color,  which  persists  for  a  day  or 
more  after  death.  These  glands  consist  of  two  elongate-oval  or  cylindrical 
glands,  of  the  opalescent  color  just  mentioned.  From  the  anterior  end 
of  each  of  these  glands  passes  the  long,  slender,  salivary  collecting  duct, 
which  unites  with  its  fellow  to  form  a  common  duct  opening  at  the  mouth 
cavity.  From  the  caudal  end  of  each  gland  is  a  flattened,  almost  hyaline, 


760  CHARLES  PAUL  ALEXANDER 

elongate   lobe  which  is  indistinctly  pseudo-segmented,   in  its  posterior 
part  being  in  intimate  connection  with  the  abundant  fat  tissue. 

Respiration  and  circulation 

The  most  important  literature  on  the  respiratory  and  circulatory  organs 
is  as  follows: 

Ptychoptera.     Grobben,  1876. 

Dicranota.     Miall,  1893:245-248. 

Phalacrocera.     Miall  and  Shelford,  1897 : 351-356. 

Liogma.     Mliggenberg,  1901. 

Ctenophora.    Anthon,  1908:551-554. 

Holorusia.     Kellogg,  1901  a;  Comstock  and  Kellogg,  1904:57-58,  60. 

Tipula  maxima.     Brown,  1910. 

Tipulinae,  Hexatomini  (especially  Tipula  paludosa  and  Limnophila  punctata).      Gerbig, 

1913. 
General  articles.     De  Meijere,  1895;  Viallanes,  1880. 

The  most  important  works  on  the  structure  of  the  tracheal  system  with 
special  reference  to  the  mechanics  of  the  spiracles,  are  those  by  Mtiggen- 
berg  (1901),  De  Meijere  (1895,  1902,  1916),  Brown  (1910),  and  Gerbig 
(1913).  Miall  and  Shelford  (1897:351-356)  discuss  in  much  detail  the 
structure  of  the  heart  in  Phalacrocera. 

In  Eriocera  spinosa,  which  may  be  considered  as  typical  for  this  group 
of  crane-flies,  there  are  two  principal  tracheal  trunks  which  lie  in  a  dorsal 
position  and  run  almost  the  length  of  the  body.  They  are  connected 
across  by  a  very  delicate,  unbranched,  simple,  tracheal  commissure,  and 
send  off  branches  laterally  to  supply  the  various  organs  of  the  body.  Near 
the  posterior  end  of  the  body  they  are  approximated,  and  just  in  front  of 
the  spiracles  they  expand  into  the  tracheal  chamber  Directly  cephalad 
of  this  chamber  the  first  lateral  branch  passes  off,  numbering  from  the 
posterior  end  forward.  Branches  2  to  8  are  large  and  much  forked. 
Just  after  leaving  the  main  tracheal  trunks,  each  of  these  sends  off  a 
ventral  branch  which  supplies  the  alimentary  canal  and  the  fat  tissue. 
Branch  3  supplies  the  region  of  the  malpighian  tubules;  branch  4,  the 
posterior  part  of  the  stomach;  branch  5,  the  anterior  part  of  the  stomach; 
branches  6  to  8,  the  esophageal  region  —  branch  6  supplying  the  pro- 
ventriculus,  branch  7  and  part  of  branch  8  the  esophagus,  and  the  remainder 
of  branch  8  the  pharyngeal  region. 

The  main  part  of  each  lateral  branch  continues  laterad,  at  its  first 
(anterior)  fork  sending  a  branch  forward  to  the  next  branch,  so  as  to 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  761 

form  a  complete  but  much-branched  lateral  trachea.  The  branches  are 
very  diffuse  and  abundant  and  the  skin  is  well  supplied.  As  already 
stated,  the  cross-commissures  are  very  delicate  and  are  unbranched  or 
nearly  so,  the  dorsal  regions  of  the  body  being  supplied  by  branches  from 
the  lateral  supply. 

The  condition  of  the  tracheae  in  Antocha,  the  only  apneustic  crane-fly 
larva  among  the  species  here  considered,  may  be  outlined  as  follows: 

The  usual  two  principal  trunks  are  present,  joined  across  on  each 
segment  by  delicate  cross-commissures  which  send  off  two  approximated 
branches  except  on  -the  eighth  segment.  Laterad  and  ventrad  of  the 
principal  trunks  are  the  delicate  lateral  tracheae.  These  are  joined  to 
the  main  trunks  by  fine  branches  inserted  about  midway  between  the 
dorsal  commissures,  toward  the  posterior  end  of  the  body  lying  nearer 
the  posterior  commissure  than  the  anterior  one. 

At  the  ninth  segment  each  trunk  sends  off  a  branch  from  its  proximal 
side,  these  branches  being  connected  by  a  long  commissure  and  supplying 
the  posterior  pair  of  gills.  The  commissure  is  about  as  long  as  that  part 
of  the  branch  between  it  and  the  trunk.  Immediately  caudad  of,  or 
just  at,  the  fork,  but  on  the  ventral  side,  a  subequal  branch  passes  into 
the  anterior  gills;  at  the  same  point  the  delicate  lateral  tracheal  trunk 
finally  ends  in  the  main  trunk.  Caudad  of  this  triple  forking  the  main 
trunk  gradually  widens  out  into  a  cylindrical  structure  which  is  truncated 
apically,  at  the  tip  passing  out  into  two  small  elongate  branches,  near 
the  base  with  about  three  delicate  branches,  two  being  dorsal  in  position 
and  one  more  lateral.  All  of  these  tracheae,  to  the  gills  and  to  the  caudal 
lobes,  send  off  many  branched  capillaries  at  frequent  intervals,  and  the 
caudal  lobes  unquestionably  function  as  tracheal  gills. 

A  comparison  of  Antocha  with  Dicranota  as  described  and  figured 
by  Miall  (1893:245-248)  shows,  in  the  latter,  distinct  spiracles  and  the 
gills  similar  but  much  smaller.  The  tracheal  arrangement  differs  in  that 
a  single  branch  on  either  side  supplies  both  gills  of  that  side,  while  the 
caudal  lobes  are  tracheated  by  a  branch  that  leaves  the  main  trunk  close 
to  the  spiracles.  (Plate  XII,  2  and  3.) 

The  arrangement  of  the  tracheae  at  the  base  of  the  wing  pad  is  described 
and  figured  for  Bittacomorpha  by  Dr.  Chapman  in  Comstock's  The 
Wings  of  Insects  (1918:36-37). 


762  CHARLES  PAUL  ALEXANDER 

The  nervous  system 
The  most  important  literature  on  the  nervous  system  is  as  follows: 

Ptychoptera.     Grobben,  1876. 

Dicranota.     Miall,  1893:241. 

Phalacrocera.     Miall  and  Shelf ord,  1897:356;  Bengtsson,  1897. 

Ctenophora.     Anthpn,  1908:556-557. 

Holorusia.     Comstock  and  Kellogg,  1904:58-59. 

In  Holorusia  the  brain,  or  supra-esophageal  ganglion,  is  composed  of 
two  principal  lobes  united  posteriorly  and  lying  above  the  esophagus. 
Beneath  the  brain  and  on  the  under  side  of  the  esophagus  lies  the  sub- 
esophageal  ganglion,  which  is  connected  with  the  anterior  end  of  the 
brain  lobe  by  the  circum-esophageal  commissures.  The  above-named 
organs  form  a  complete  ring,  or  collar,  around  the  alimentary  canal.  The 
ventral  nervous  system  leads  backward  from  the  subesophageal  ganglion 
on  the  ventral  surface  of  the  body  wall.  In  the  thorax  there  are  four 
closely  approximated  ganglia  representing  the  thoracic  and  the  first 
abdominal  segments.  Beyond  these  and  located  in  the  abdomen  are 
six  abdominal  ganglia.  The  ganglia  send  off  four  large  nerve  trunks. 

The  condition  is  similar  in  Ctenophora  and  in  Tipula,  there  being  seven 
abdominal  ganglia,  of  which  the  first  is  located  in  the  thorax  and  the  last 
six  in  the  abdomen.  In  Phalacrocera  there  are  eight  abdominal  ganglia, 
the  first  being  usually  found  in  the  first  abdominal  segment  and  the 
eighth  in  segment  10.  The  head  ganglia  are  all  independent  of  one 
another,  not  fused  into  a  complex.  The  longitudinal  commissures  between 
the  first  five  ventral  ganglia  are  double,  and  those  between  the  eight 
abdominal  ganglia  are  simple  (Bengtsson).  In  Dicranota  there  are  like- 
wise eight  abdominal  ganglia,  the  first  being  located  in  the  first  abdominal 
segment  and  the  others  in  segments  5  to  10,  there  being  two  close  together 
in  the  tenth  segment. 

The  reproductive  system 
The  most  important  literature  on  the  reproductive  system  is  as  follows: 

Ptychoptera.     Grobben,  1876. 

Dicranota.     Miall,  1893:248. 

Phalacrocera.     Miall  and  Shelf  ord,  1897:356-357. 

Holorusia.     Comstock  and  Kellogg,  1904:58. 

The  reproductive  organs  appear  early  in  the  larval  development.  In 
Dicranota  the  testes  lie  in  the  ninth  segment  and  form  elongate  capsules, 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  763 

when  sufficiently  advanced  giving  off  the  vasa  deferentia  from  the  inner 
side  of  the  hinder  end.  Even  in  larvae  not  fully  matured,  Miall  found 
ripe  spermatazoa  mixed  with  spermatoblasts.  No  division  of  the  testes 
into  follicles  is  apparent.  In  Phalacrocera,  also,  ripe  spermatazoa  may 
be  found  in  the  older  larvae. 

In  Phalacrocera  the  ovaries  form  a  pair  of  cylindrical  bodies  tapering 
to  each  end,  lying  on  either  side  of  the  beginning  of  the  intestine.  The 
ovaries  are  very  peculiar  in  structure,  seeming  to  be  adapted  to  the  almost 
simultaneous  discharge  of  all  the  eggs. 

The  muscular  system 
The  most  important  literature  on  the  muscular  system  is  as  follows: 

Body  musculation 

Ptychoptera.     Grobben,  1876. 

Dicranota.     Miall,  1893:241. 

Ctenophora.     Anthon,  1908:545. 

Holorusia.     Comstock  and  Kellogg,  1904:59-60. 

Musculation  of  the  mouth  parts 
Phalacrocera.     Bengtsson,  1897. 

The  fat  bodies,  or  adipose  tissue 
The  most  important  literature  on  the  adipose  tissue  is  as  follows: 

Dicranota.  M#all,  1893:241-242. 
Ctenophora.  Anthon,  1908:554. 
Holorusia.  Comstock  and  Kellogg,  1904:56. 

The  imaginal  disks,  or  histoblasts 
The  most  important  literature  on  the  imaginal  disks  is  as  follows: 

Phalacrocera.     M  iall  and  Shelf ord,  1897 : 357-358. 

Ctenophora.     Anthon,  1908:555. 

Holorusia.     Kellogg,  1901  b;  Comstock  and  Kellogg,  1904:61. 

PHYLOGENY 

The  origin  and  phylogenetic  development  of  the  various  groups  of 
crane-flies  is  still  largely  problematical.  The  evidence  supplied  by  the 
adult  flies  of  the  most  generalized  living  groups  points  to  an  ancestor 
which  in  many  respects  resembled  the  recent  Mecoptera,  or  scorpion 


764  CHARLES  PAUL  ALEXANDER 

flies.  That  this  resemblance  is  presumably  not  fundamental  is  suggested 
by  the  immature  stages  of  these  same  generalized  forms,  which  show 
but  few  features  that  can  be  reconciled  with  those  of  a  panorpid-like 
ancestor.  Unfortunately  the  fossil  record  helps  but  little.  At  the  begin- 
ning of  the  Tertiaries,  crane-flies  were  already  numerous  both  in  species 
and  in  individuals,  but  in  most  cases  they  are  clearly  referable  to  existing 
genera.  It  is  evident,  then,  that  the  investigator  must  go  still  further 
back  for  his  evidence,  and  the  fossil  crane-flies  of  the  Upper  Mesozoic 
are,  unfortunately,  still  largely  unavailable.  The  phylogenetic  develop- 
ment of  the  group  may  be  considered  from  two  standpoints,  the  geological 
record  and  comparative  morphology. 

THE    GEOLOGICAL   RECORD 

The  most  important  works  on  fossil  crane-flies  are  those  of  Loew 
(1850),  Scudder  (1894),  Handlirsch  (1906-08,  and  1910,  a  and  b),  Meunier 
(1906),  and  Cockerell  (1910,  1913,  1916,  and  1917,  a  and  b).  The  writer 
is  indebted  to  Professor  Cockerell  for  assistance  in  determining  the  age 
of  many  of  the  fossil-bearing  beds.  Excellent  accounts  of  the  various 
deposits  may  be  found  in  Cockerell  (1913)  and  in  Tilly ard  (1917). 

The  Tipuloidea,  representing  the  superfamily  of  crane-flies,  is  herein 
considered  as  being  constituted  of  six  families,  of  which  two,  the 
Eoptychopteridae  and  the  Architipulidae,  did  not  survive  the  Mesozoic 
period.  The  Eoptychopteridae  are  known  only  from  the  Mecklenburg 
Lias  (lowest  Jurassic),  and  include  but  three  monotypic  genera.  The 
Architipulidae  are  known  from  the  same  formation  and  include  eight 
species  arranged  in  three  genera.  The  other  four  families  contain  recent 
forms  and  are  included  in  this  paper. 

The  North  American  Eocene  and  Miocene,  as  represented  by  the  White 
River  and  Green  River  beds  and  the  Florissant  shales,  respectively,  give 
evidence  of  having  had  a  northern  fauna,  especially  in  the  Eocene.  This 
is  well  shown  by  the  great  development  of  the  Cylindrotominae,  which 
in  the  White  River  basin  almost  dominated  the  crane-fly  fauna  during 
the  Eocene.  It  seems  probable,  moreover,  that  the  group  was  forced 
into  colder  regions  of  the  globe  during  the  Oligocene,  when  the  tropical 
element  reached  far  to  the  north.  No  group  of  crane-flies  that  can  be 
considered  as  being  tropical  has  yet  been  found  in  the  Florissant.  On  the 
other  hand,  the  European  Oligocene,  as  shown  by  the  Gurnet  Bay  beds 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  765 

and  the  lower  Oligocene  Baltic  amber,  has  a  considerable  tropical  element 
apparent  —  as,  for  instance,  Styringomyia,  found  in  both  formations, 
and  Trentepohlia,  Polymera,  and  Brachypremna,  the  last  two  being 
amber  records.  In  connection  with  these  two  last-named  genera,  as 
fossils  known  only  from  the  Baltic  amber,  it  should  be  noted  that  each 
is  found  living  to-day  only  in  America,  where  they  are  confined  to  the 
austral  and  tropical  regions. 

The  Tanyderidae  are  represented  by  the  Oligocene  Macrochile  (amber) 
and  Etoptychoptera.  At  present  this  is  still  a  very  small  group,  including 
but  two  genera  and  nine  species. 

The  Ptychopteridae  include  a  Ptychoptera  from  the  Bohemian  Krot- 
tensee  (probably  upper  Oligocene)  and  a  Bittacomorpha  from  the 
Florissant. 

The  Rhyphidae  (Trichocerinae)  are  represented  by  two  species  from 
the  Baltic  amber  and  one  or  two  additional  species  from  the  Miocene. 

The  Tipulidae  may  be  considered  in  general  terms  only.  But  one 
tipulid  is  listed  by  Handlirsch  as  being  known  from  the  Mesozoic.  In 
the  earliest  Tertiaries,  however,  a  variety  of  species  is  found  in  this  family 
which  almost  parallels  their  recent  development. 

The  Limnobiini  first  appeared  in  the  Eocene  (Dicranomyia,  Gera- 
nomyia).  In  the  lower  Oligocene,  numerous  species  of  Dicranomyia  and 
Rhipidia  occurred.  In  the  Miocene,  Dicranomyia  was  common,  tho 
probably  not  so  rich  in  species  as  at  present.  Rhamphidia  is  found  in 
amber,  and  Antocha  has  been  described  from  the  Florissant  Miocene  by 
Scudder;  the  latter  record,  however,  seems  very  doubtful  to  the  writer, 
judging  from  Scudder's  figure  and  description. 

The  Hexatomini  are  represented  by  Ula,  Limnophila,  Eriocera,  and 
Polymera  in  amber,  and  by  Limnophila  in  the  Florissant  shales. 

There  is  no  record  of  the  Pediciini  being  preserved  as  fossils. 

The  Eriopterini  are  well  represented  in  the  Baltic  amber  by  Erioptera, 
Gnophomyia,  Gonomyia,  and  related  genera,  and  also  in  the  Miocene  by 
Gonomyia,  Cladura,  and  others.  Toxorhina,  an  apparent  relative  of 
Elephantomyia,  occurs  in  the  Baltic  amber. 

The  Styringomyiini  are  represented  by  the  only  living  genus,  Styringo- 
myia, a  fly  of  uncertain  affinities,  in  the  Gurnet  Bay  Oligocene  (Cockerell, 
1917  c  and  1919)  and  in  amber,  reappearing  in  the  Pleistocene  African 
copal. 


766  CHARLES  PAUL  ALEXANDER 

The  dominance  of  the  Cylindrotominae  (genus  Cyttaromyia)  in  the 
Eocene  and  Miocene  of  the  North  American  fauna  has  already  been 
mentioned.  No  records  of  this  group  from  the  European  Oligocene 
are  available.  The  recent  species  of  the  subfamily  are  practically  all 
forms  belonging  to  cold,  temperate  regions,  the  few  Oriental  species  of 
Stibadocera  coming  from  mountains  at  considerable  altitudes. 

The  Tipulinae  have  been  found  as  far  back  as  the  Mesozoic,  but  the 
records  are  not  entirely  satisfactory.  In  the  lowermost  Tertiaries,  how- 
ever, undoubted  tipuline  forms  occur.  Species  occur  in  the  Green  River 
shales  of  Colorado  (Eocene).  The  group  was  well  represented  in  the 
Oligocene  (Baltic  amber,  Tulameen  beds  of  British  Columbia,  Krottensee, 
and  Gurnet  Bay),  and  was  very  common  in  the  Miocene  (Radoboj,  and 
especially  in  the  Florissant  of  Colorado,  where  some  twenty-five  species 
of  Tipula  and  closely  allied  genera  or  subgenera  have  been  described  by 
Scudder  and  Cockerell). 

Tipulidae  of  the  Pleistocene  are  not  numerous,  only  a  few  having  been 
made  known  from  the  refuse  of  lake  dwellings  in  England  (Dicaera, 
apparently  related  to  Ctenophora),  and  in  the  African  copal,  including 
such  genera  as  Styringomyia,  Elephantomyia,  and  Toxorhina. 

COMPARATIVE   MORPHOLOGY 

The  morphology  of  the  various  stages  of  crane-flies  has  been  detailed 
elsewhere  in  this  paper  and  need  not  be  repeated  here. 

PHYLOGENETIC  CONSIDERATIONS 

The  eucephalous  familes  of  crane-flies  are  undoubtedly  lower,  phylo- 
genetically,  than  the  Tipulidae,  and  the  latter  have  been  derived  from 
the  former.  The  generalized  type  recurs  in  all  three  subfamilies  of  the 
Tipulidae,  and  it  is  uncertain  which  of  these  three  should  be  placed 
lowermost.  Presumably  all  three  groups  arose  from  an  immediate  com- 
mon ancestor,  or  the  Tipulinae  and  the  Limnobiinae  arose  from  one  point  of 
the  tree,  the  Cylindrotominae  developing  from  the  limnobiine  stem  at  a 
somewhat  later  period.  The  accompanying  phylogenetic  tree  (Plate  XII ,  4) 
graphically  illustrates  this  apparent  evolution  of  the  group.  The  Limno- 
biini  show  but  little  deviation  from  the  fundamental  type.  From  the 
level  of  the  lowermost  Hexatomini  (Ula  and  Epiphragma),  in  close 
proximity  to  the  Limnobiini,  the  remaining  groups  of  crane-flies  can  be 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  767 

derived  —  the  Pediciini  and  the  Eriopterini  on  the  one  hand,  the  hexato- 
mine  divisions  on  the  other.  The  highest  levels  of  development  of  their 
respective  types  are  apparently  Diotrepha  (Limnobiini),  Dicranota 
(Pediciini),  Gonomyia  (Eriopterini),  and  Hexatoma  (Hexatomini). 

The  immature  stages  of  the  Tipulidae  of  the  antipodal  regions  (Australia 
and  Chile)  are  entirely  unknown,  and  their  discovery  may  bring  to  light 
striking  novelties  that  may  well  upset  the  present  ideas  of  classification. 
The  largest  remaining  gaps  in  the  present  knowledge  of  the  Nearctic 
fauna  relate  to  the  genera  Atarba,  Toxorhina,  Cryptolabis,  Phyllolabis, . 
and  Megistocera,  but  it  is  not  expected  that  any  major  groups  will  have 
to  be  created  for  their  reception. 

LIFE  HISTORIES   OF  CRANE-FLIES,   WITH  KEYS  AND  DESCRIPTIONS  OF 

THE  SPECIES 

Various  classifications  and  arrangements  of  Diptera  have  been  proposed 
in  the  past,  and  the  principal  evolutionary  changes  in  the  entomologist's 
conception  of  classification  are  herein  indicated. 

The  groupings  in  the  superfamily  Tipuloidea  have  been  very  diverse. 
Brauer  (1883)  separated  the  eucephalous  families  from  the  Tipulidae 
(Polyneura).  Osten  Sacken  (1893)  placed  the  Tipulidae  in  the  Nemocera 
vera,  the  Rhyphidae  in  the  Nemocera  anomala.  Coquillett  (in  Howard, 
1912:85-86)  included  the  Tipulidae  in  his  superfamily  Tipuloidea,  but 
separated  the  Rhyphidae,  placing  the  latter  in  Bibionoidea.  Lameere  (1906) 
gave  a  classification  that  had  little  basis  in  fact,  the  Rhyphidae  being 
considered  by  him  as  Brachycera  and  the  Ptychopteridae  being  placed 
with  the  Culicidae.  Knab  (1915),  working  on  the  respiratory  systems 
of  the  larvae,  demonstrated  the  hitherto  unnatural  groupings  of  the 
families  of  the  Nemocera,  and  arranged  the  crane-fly  families,  along 
with  a  few  others,  in  the  group  Polyneura,  a  group  coordinate  in  rank 
with  the  previous  conception  of  the  Nematocera.  The  latest  grouping, 
that  of  Malloch  (191 5- 17  b),  was  based  on  a  broad  knowledge  of  both  the 
adult  flies  and  the  immature  stages;  and  here,  for  the  first  time,  one  gets 
the  true  conception  of  the  superfamily  limits  —  the  inclusion  of  the 
Ptychopteridae,  the  Tipulidae,  and  the  Rhyphidae  in  a  single  major  group. 
Malloch's  arrangement  is  herein  adopted,  with  the  following  exceptions: 
(1)  the  Limnobiidae  are  not  held  as  constituting  a  separate  family,  but 
are  united  with  the  Tipulidae;  (2)  the  family  Tanyderidae  has  been 


768  CHARLES  PAUL  ALEXANDER 

erected,  to  include  the  generalized  Tanyderina  which  have  hitherto  been 
placed  with  the  Ptychopteridae. 

The  immature  stages  of  the  four  families  of  the  Tipuloidea  are  readily 
separable.  The  larvae  of  the  Tipulidae  can  be  confused  only  with  those 
of  certain  low  brachycerous  forms,  as,  for  example,  the  Leptidae.  In  the 
Brachycera  the  mandibles  work  vertically  and  parallel  to  each  other; 
in  the  Nematocera,  including  the  Tipuloidea,  they  operate  transversely 
or  obliquely  against  the  teeth  of  the  mentum  and  the  hypopharynx. 
The  presence  of  fleshy  lobes  surrounding  the  spiracular  disk  is  a  character 
possessed  by  almost  all  Tipulidae.  The  larvae  of  a  few  groups  of  brachyc- 
erous Diptera,  such  as  certain  Leptidae,  Sciomyzidae,  and  other  families, 
possess  entirely  similar  caudal  lobes  but  are  readily  recognized  by  the 
small  and  very  reduced  head  capsule. 

The  eucephalous  families  of  the  Tipuloidea  may  be  distinguished  by 
means  of  the  characters  indicated  in  the  following  keys: 

Larvae 

1.  Body  eucephalous,  head  non-retractile;  amphipneustic  or  metapneustic 2 

Head  incomplete  behind,  retractile;  not  amphipneustic TIPULIDAE  (p.  791) 

2.  Caudal  end  of  body  prolonged  into  a  slender  breathing  tube;  metapneustic 3 

Caudal  end  of  body  not  prolonged  into  a  breathing  tube;  amphipneustic. 

RHYPHIDAE  (p.  787) 

3.  Breathing  tube  stouter,  non-retractile;  gills  large,  pinnately  branched;  punctures  of  head 

multisetose;  found  in  wet  decaying  wood TANYDERIDAE,  supp.  (p.  769) 

Breathing    tube    slender,  completely    or    partly    retractile;   gills    slender,     cylindrical, 
unbranched;  punctures  of  head  with  simple  or  plumose  hairs;  found  in  wet  earth. 

PTYCHOPTERIDAE  (p.  772) 

Pupae 

1.  One  of  the  pronotal  breathing  horns  greatly  elongated,  much  longer  than  the  body,  the 

other  breathing  horn  very  short,  abortive.     (Family  PTYCHOPTERIDAE,  p.  772) 2 

Breathing  horns  short,  or,  if  elongated   (some   Tipulinae),  not  longer  than  the  body 
and  the  difference  in  size  not  so  apparent 3 

2.  Tarsal  sheaths  lying  side  by  side,  parallel Ptychopterinae  (p.  773) 

The  fore  tarsal  sheaths  overlying  the  middle  pair,  the  four  middle  and  hind  tarsi  the 

longest,  parallel Bittacomorphinae  (p.  779) 

3.  Tarsal  sheaths  overlying  one  another  in  pairs RHYPHIDAE  (p.  787) 

Tarsal  sheaths  lying  side  by  side TIPULIDAE  (p.  791) 

The  pupae  of  the  Tar^deridae  are  still  unknown. 

It  is  possible  that  Bittacomorphella  (page  779)  has  short  breathing 
horns;  in  this  case  this  genus  would  run  down  to  couplet  3  above,  but 
by  the  arrangement  of  the  tarsal  sheaths  it  runs  out  as  indicated  in 
couplet  2. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  769 

FAMILY  Tanyderidae 

Larva  (supposition). —  Body  eucephalous,  metapneustic.  Integument  smooth,  shiny  white. 
Last  two  segments  of  abdomen  produced  into  a  stout,  non-retractile  breathing  tube,  which 
is  distinctly  five-lobed  at  the  tip.  Tracheal  gills  two,  very  large,  pinnately  branched.  Head 
with  scattered  punctures,  which  are  multisetose;  lateral  plates  of  head  united  across  venter. 
Mandibles  opposed,  narrow,  tridentate.  Maxilla  with  palpus  two-segmented,  the  outer 
lobe  digitiform,  pubescent.  Labrum  small,  semi-circular,  with  two  punctures;  clypeus  with 
four  punctures. 

Pupa.- —  Unknown. 

The  Tanyderidae  comprise  a  group  of  primitive  crane-flies  including 
but  three  recent  genera,  with  ten  species.  Of  these  the  genus  herein 
considered,  Protoplasa,  with  three  known  species,  is  found  in  the  Northern 
Hemisphere.  All  that  is  known  concerning  the  biology  of  supposed 
species  of  this  group  pertains  to  Protoplasa  fitchii  and  is  discussed  below. 

Genus  Protoplasa  Osten  Sacken  (Gr.  first  +  to  form) 

1859     Protoplasa  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  252. 
1878    Idioplasta  0.  S.     Cat.  Dipt.  N.  Amer.,  p.  222. 

The  genus  Protoplasa  is  confined  to  temperate  North  America.  There 
are  three  known  species,  of  which  P.  vipio  O.  S.  and  P.  vanduzeei  Alex, 
are  western  in  their  distribution,  while  P.  fitchii  O.  S.  is  eastern.  The 
general  characters  of  the  supposed  larva  are  given  above;  the  detailed 
account  in  connection  with  the  species  P.  fitchii  follows. 

Protoplasa  fitchii  O.  S. 
1859  Protoplasa  fitchii  0.  S.    Proc.  Acad.  Nat.  Sci.  Phila.,  p.  252. 

The  remarkable  insect  Protoplasa  fitchii  is  one  of  the  rarest  of  the 
local  crane-flies.  There  are  scarcely  a  score  of  specimens  extant  in  the 
various  collections  of  the  country,  most  of  which  are  from  the  mountainous 
section  of  North  Carolina.  The  fly  has  not  been  reared,  but  the  writer  has 
in  his  possession  a  larva  that  he  refers  with  much  confidence  to  this  species. 
It  is  one  of  the  most  remarkable  dipterous  larvae  that  have  ever  come  to 
the  writer's  notice,  and,  whether  or  not  it  belongs  to  Protoplasa,  it  should 
certainly  be  called  to  the  attention  of  entomologists  in  the  hope  that  it 
may  some  day  be  bred  and  its  identity  confirmed  or  ascertained.  These 
peculiar  dipterous  larvae  were  discovered  by  H.  S.  Barber,  C.  T.  Greene,  and 


770  CHARLES  PAUL  ALEXANDER 

R.  C.  Shannon,  on  the  Potomac  River  near  the  mouth  of  Dead  Run,  Fair- 
fax County,  Virginia.  They  were  found  during  the  latter  part  of  May,  1916, 
in  a  much-decayed  maple  log,  where  they  were  associated  with  the  larvae 
of  the  syrphid  fly,  Temnostoma  bombylans  (Fabr.)  (Barber,  1913).  Greene 
attempted  to  rear  the  larvae  procured  at  this  time,  but  did  not  succeed. 
In  May,  1917,  he  sent  the  writer  one  of  the  preserved  larvae  for  study. 
Later  Dr.  Viereck  was  interested  in  the  matter,  and  on  May  27  he  procured 
one  fine,  healthy  larva,  which  was  sent  to  the  writer  at  Ithaca,  New  York. 
It  arrived  safely  on  the  29th,  and  was  at  once  placed  in  rearing.  Unfortu- 
nately this  larva  died  the  day  after  it  came  into  the  writer's  possession, 
and  this  remarkable  insect  still  remains  unreared.  Associated  with  this 
larva  in  the  pieces  of  decaying  maple  in  which  it  was  shipped,  were 
larvae  of  Temnostoma  and  a  larva  and  a  pup"a  of  the  tipulid  Epiphragma 
solatrix. 

The  evidences  that  this  larva  is  that  of  Protoplasa  are  numerous. 
It  belongs,  without  doubt,  to  the  Nematocera,  and  the  immature  stages 
of  all  the  remaining  families  of  that  division  have  been  made  known. 
Many  features  in  this  larva  point  strongly  to  the  condition  in  other  families 
of  crane-flies.  The  eucephalous  condition  of  the  head,  and  the  elongate 
breathing  tube,  are  suggestive  of  the  Ptychopteridae;  the  five-lobed 
spiracular  disk,  the  anal  tracheal  gills,  the  metapneustic  respiratory 
system,  and  other  features,  are  very  similar  to  conditions  in  certain 
Tipulidae.  However,  there  are  conditions  obtaining  here  that  are  found 
nowhere  else  in  the  Diptera,  so  far  as  is  known  to  the  writer,  such  as  the 
combination  of  a  eucephalous  head  of  primitive  organization,  a  stout, 
non-retractile  breathing  tube,  the  large,  pinnately  branched  anal  gills, 
the  multisetose  punctures  on  the  head,  and  the  details  of  structure  of  the 
mouth  parts.  The  multisetose  punctures  are  suggestive  of  the  branched 
or  plumose  hairs  of  Ptychoptera,  and  give  a  possible  hint  of  the  origin 
and  ancestry  of  this  condition  in  the  latter  group.  The  writer  cannot 
but  regard  it  as  suggestive  that  the  larvae  are  found  in  direct  association 
with  those  of  Epiphragma  in  saturated  decaying  wood.  He  has  mentioned 
in  other  papers  (Alexander,  1910:254,  and  1919  d:  883,  915)  the  remarkable 
superficial  resemblance  that  exists  between  the  adult  flies  of  Epiphragma 
fasdpennis  and  those  of  Protoplasa,  both  forms  having  very  hand- 
somely banded  wings  of  a  pattern  not  found  in  other  species  in  the  local 
fauna. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  771 

The  only  observations  on  the  adult  flies  that  the  writer  has  ever  been 
able  to  make  were  in  reference  to  five  specimens  taken  in  New  York 
State  in  1909  (Alexander,  1910:254).  These  were  swept  from  rich  vege- 
tation along  the  banks  of  the  Sacandaga  River  in  northeastern  New  York. 
The  adult  flies  have  never  been  recorded  from  the  vicinity  of  Washington, 
but  are  regional,  since  the  species  ranges  thruout  the  eastern  United 
States  as  far  south  as  Georgia  (in  the  mountains). 

Larva. — Total  length,  11.8  mm. 

Length  excluding  breathing  tube,  5.6-6  mm. 

Length  of  breathing  tube  alone,  5.8-6.2  mm. 

Length  of  ninth  segment  of  abdomen  (chitinized  part  of  breathing  tube),  3.4-3.6  mm. 

Length  of  anal  gills,  2-2.1  mm. 

Diameter  of  body,  2.3  mm. 

Coloration  pure  white,  head  and  apical  two-thirds  of  breathing  tube  pale  brown,  chitin- 
ized. In  the  living  larva  there  is  a  transparent,  subcircular  area  on  the  dorsum  of  segment  7 
thru  which  the  beating  of  the  heart  can  be  clearly  seen,  but  in  preserved  specimens  this  area 
is  hard  to  perceive.  (Plate  XIII,  5  ) 

Body  eucephalous,  entirely  smooth,  shiny,  dorsum  of  segments  2  to  9  each  with  a  trans- 
verse group  of  tiny  points.  Thorax  with  the  first  segment  longer  than  the  succeeding  two 
segments  taken  together,  narrowed  anteriorly,  broader  behind;  mesothoracic  segment  about 
half  as  long  as  the  first  and  about  twice  as  long  as  the  very  narrow  metathorax.  Abdominal 
segments  gradually  increasing  in  length  to  the  seventh;  segments  8  and  9  abruptly  narrowed 
into  a  stout,  elongate  breathing  tube  which  is  about  equal  in  length  to  remainder  of  body; 
basal,  or  proximal,  part  of  tube  (segment  8)  shorter  than  terminal  part  (segment  9)  and 
not  chitinized;  segment  9  elongate,  chitinized,  with  a  deep  transverse  constriction  a  short 
distance  before  tip;  this  apical  part,  and  the  region  just  before  the  constriction,  smooth, 
the  remainder  of  the  last  segment  with  numerous  delicate  transverse  wrinkles.  Spiracular 
disk  (Plate  XIII,  6)  surrounded  by  five  lobes,  one  dorsal,  two  lateral,  and  two  ventral 
in  position;  dorsal  lobe  the  smallest,  lateral  lobes  the  broadest;  lobes  tipped  with  sharp, 
chitinized  spines,  which  are  continued  for  a  short  distance  down  the  outer  face  of  each  lobe; 
fringes  of  long  hairs  along  margins  of  lobes;  disk  with  a  brownish  tinge  around  spiracles 
and  an  elongate-oval  median  mark  between,  and  slightly  below,  the  spiracular  level;  inner 
face  of  ventral  lobes  with  indistinct,  tiny,  brown  punctures.  Spiracles  large,  separated  by 
a  distance  about  equal  to  diameter  of  one.  Anal  tracheal  gills  two,  one  on  either  side  of 
the  base  of  segment  8,  very  large  and  pinnately  branched,  there  being  about  fifteen  branches 
to  each  gill;  each  of  these  branches  constricted  into  four  or  five  lobules;  lobes  at  their  tips 
broadly  obtuse  (Plate  XIII,  9). 

Head  short  and  broad,  epicranium  chitinized.  Across  ventral  face,  the  sclerite  firmly 
united  by  a  narrow  transverse  band;  median  part  densely  punctulate,  this  area  delimited 
laterally  and  posteriorly  by  an  indistinct  curved  line.  On  dorsum,  the  prefrons  conspicuous 
and  the  clypeus  distinct  from  the  labrum.  Head  and  mouth  parts  provided  with  numerous 
setiferous  punctures,  each  of  which  is  multisetose  (from  five  to  eight  setae  to  each  puncture). 
Chaetotaxy  of  epicranium  (Plate  XIII,  7)  with  three  lateral  groups  of  long  bristles,  the 


772  CHARLES  PAUL  ALEXANDER 

posterior  group  being  the  longest  and  most  dorsal  in  position;  on  ventral  face,  four  punctures 
on  either  side  (Plate  XIII,  8);prefrontal  sclerite  apparently  lacking  setae,  but  on  the  epicranium 
proper,  along  margin  of  prefrons,  a  longitudinal  group  of  six  punctures  on  either  side,  the 
anterior  pair  the  longest.  Labrum  semicircular  in  outline,  outer  margin  fringed  with  long, 
stiff  hairs;  dorsal  surface  with  two  transverse  punctures.  Clypeus  narrowly  transverse, 
with  four  setiferous  punctures  in  a  transverse  row.  Dorsal  median  part  of  epicranium 
forming  a  subquadrate  lobe  whose  lateral  angles  bear  brushes  of  hairs;  laterad  of  these  angles 
a  rounded  hollow  overlapped  by  a  ^at,  bilobed  operculum  (this  may  be  some  modification 
of  the  antennae  which  should  occupy  about  this  position  on  the  head).  Mandible  elongate, 
strongly  chitinized,  tridentate,  the  apical  tooth  the  longest  and  more  acute  than  the  others, 
the  middle  tooth  bluntly  obtuse;  inner  posterior  angle  of  mandible  produced  proximad 
beneath  outer  lobe  of  maxilla  into  a  bluntly  rounded  lobe;  mandible  with  a  group  of  curved 
setae  on  scrobe  near  base,  and  a  group  of  longer  bristles  lying  distad  and  mesad  of  these 
setae  on  dorsal  face  of  mandible.  Maxilla  with  base,  in  a  position  of  rest,  concealed  beneath 
head-chitin;  palpus  stout,  antenniform,  two-segmented,  the  basal  segment  short  and  stout, 
the  apical  segment  much  shorter  and  narrower,  with  a  small,  lateral,  sensory  papilla;  outer 
lobe  of  maxilla  produced  cepha-lad  as  a  stout,  digitiform  lobe  which  is  densely  hairy. 

FAMILY  Ptychopteridae 

Larva. —  Body  eucephalous,  metapneustic,  long  and  slender,  the  caudal  end  prolonged 
into  a  more  or  less  completely  retractile  breathing  tube  bearing  the  spiracles  at  the  tip. 
Lobes  surrounding  spiracular  disk  indistinct.  Anal  gills  two,  elongate-cylindrical, 
unbranched.  Integument  with  tiny  hairs  (Ptychoptera),  or  with  slight  warty  pro- 
tuberances (Bittacomorpha),  or  with  conspicuous  elongate  tubercles  (Bittacomorphella). 
Pseudopods  on  abdominal  segments  1  to  3  each  bearing  a  curved  claw.  Head  complete; 
eyespots  distinct.  Mandibles  opposed.  Mentum  many-toothed  (Ptychopterinae)  or  merely 
bilobed  (Bittacomorphinae). 

Pupa.' —  Usually  with  one  of  the  two  pronotal  breathing  horns  greatly  elongated,  much 
longer  than  the  body  (this  may  not  be  true  in  Bittacomorphella);  in  Ptychoptera  and 
Bittacomorpha  it  is  the  right  horn  that  is  elongated,  the  left  being  degenerated;  in  Bitta- 
comorphella the  right  horn  is  degenerated.  Tarsal  sheaths  all  parallel  in  the  Ptychopterinae, 
the  fore  pair  overlying  the  middle  pair  in  the  Bittacomorphinae.  Abdomen  covered  with 
setiferous  tubercles  arranged  in  transverse  rows  on  tergites  and  sternites,  and  more  or  less 
in  longitudinal  rows  on  pleurites.  Cauda  with  a  powerful  dorsal  median  lobe  near  base  of 
segment  8. 

The  family  Ptychopteridae  includes  three  genera  falling  in  two  tribes: 
the  Ptychopterinae  including  the  single  genus  Ptychoptera,  and  the 
Bittacomorphinae  including  the  "  phantom  crane-flies,"  Bittacomorpha 
and  Bittacomorphella.  The  habits  of  these  species  are  discussed  below 
in  connection  with  the  various  genera.  The  following  keys  separate 
the  subfamilies  of  the  Ptychopteridae: 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  773 

Larvae 

Mentum  with  outer  margin  finely  serrated;  mandibles  with  three  large  outer  teeth;  pseudo- 
pods  small;  coloration  yellow  or  brown Ptychopterinae  (p.  773) 

Mentum  bilobed,  not  toothed;  mandibles  with  a  single  large  outer  tooth;  pseudopods  prom- 
inent, each  with  a  conspicuous  curved  claw;  coloration  rusty  red  or  black. 

Bittacomorphinae  (p.  779) 
Pupae 

All  tarsi  lying  parallel;  wing  pads  with  M  branched Ptychopterinae  (p.  773) 

Fore  tarsi  lying  above  middle  tarsi;  wing  pads  with  M  unbranched. 

Bittacomorphinae  (p.  779) 

The  immature  stages  of  this  group  of  flies  have  long  been  known,  having 
been  investigated  by  Reaumur  (1740),  Lyonet  (1832),  and  other  early 
workers.  The  immature  stages  of  the  common  American  species 
Pty  chapter  a  rufocincta  are  herein  recorded  for  the  first  time.  The  unknown 
Ptychoptera  described  by  Malloch  (1915-17  b:  240-241)  is  evidently  P. 
lenis  O.  S.  and  is  here  considered  as  that  species. 

The  most  important  literature  on  the  Ptychopteridae  is  as  follows: 

Ptychoptera  paludosa Larva,  pupa,  general. .  Reaumur,  1740,  pi.  6. 

Ptychoptera  paludosa General Wesenberg-Lund,  1915:348-351. 

Ptychoptera  lacustris Larva Beling,  1886: 171-172. 

Ptychoptera  contaminata Larva,  general Van  Gehuchten,  1890.  (Histol- 
ogy of  the  alimentary  canal.) 

Ptychoptera  albimana General Cameron,  1917:65. 

Ptychoptera  albimana Larva,  pupa,  general. .  Topsent,  1914-16. 

Ptychoptera  sp.  (lenis  supp.) Larva Malloch,  1915-17 b: 240-241. 

Ptychoptera  sp General Leger  and  Duboscq,  1909.  (Pro- 
tozoan parasites.) 

Ptychoptera  sp Larva,  general Grobben,  1876.     (Morphology.) 

Ptychoptera  sp Larva,  general De  Meijere,  1916: 188-191,  figs. 

14-20.  (External  morphology.) 

Ptychoptera  sp General Lyonet,  1832. 

Bittacomorpha  clavipes Larva,  pupa,  general. .  Hart,  1898  [1895]:  189-195. 

Bittacomorpha  clavipes. General Needham  and  Betten,  1901:574. 

Bittacomorpha  clavipes General Howard,  1912:95-96. 

Bittacomorpha  clavipes Larva,  pupa Malloch,  1915-17b: 239-240. 

Bittacomorpha  clavipes General Weston  and  Turner,  1917:53. 

Van  der  Wulp  (1857),  Miall  (1895),  Grunberg  (1910),  and  Brunetti 
(1912)  also  give  general  accounts  of  the  immature  stages  of  Ptychoptera, 
for  the  most  part  taken  from  the  earlier  works  cited  above. 

SUBFAMILY  Ptychopterinae 
Genus  Ptychoptera  Meigen  (Gr.  fold  +  wing) 

1800    Liriope  Meig.     Nouv.  Class.  Mouch.,  p.  14  (nomen  nudum). 
1803    Ptychoptera  Meig.     Illiger's  Mag.,  vol.  2,  p.  262. 
1856    Ctenoceria  Rond.     Dipt.  Ital.  Prodr.,  vol.  1,  p.  187. 


774  CHAKLES  PAUL  ALEXANDER 


.-  —  Head,  oval  to  subpyriform,  broadest  behind,  narrowed  in  front,  not  heavily 
patterned  above;  setae  of  head  branched  or  plumose.  Submentum  large,  usually  but  a  little 
narrowed  posteriorly,  the  anterior  lateral  angles  slightly  produced;  mentum  broader  than 
long,  outer  margin  with  numerous  teeth  (18-22).  Prementum  with  palpi  rather  small, 
lying  parallel,  densely  hairy  on  outer,  or  ventral,  face.  Mandible  with  three  powerful  outer 
teeth  and  about  six  smaller  inner  teeth.  Maxilla  with  the  cardines  elongate  -triangular, 
with  three  setiferous  punctures;  maxillary  palpi  antenniform,  cylindrical.  Labrum  broad, 
transverse,  with  dense  tufts  of  hair  beneath.  Integument  almost  smooth,  without  prominent 
warty  elevations.  Pseudopods  low,  with  small  claws. 

Pupa.  —  Head  small,  with  a  reduced  cephalic  crest.  Sheaths  of  maxillary  palpi  elongate, 
the  tips  projecting  around  knee  joints  of  fore  legs.  Leg  sheaths  with  all  the  tarsi  parallel, 
not  overlapping  one  another.  Wing  sheaths  with  media  branched.  Pronotal  breathing 
horns  very  unequal,  one  (usually  the  right)  exceedingly  elongate,  longer  than  remainder 
of  body,  the  other  tiny,  degenerate.  Abdomen  set  with  transverse  and  longitudinal  rows 
of  small  setiferous  tubercles  on  the  segments. 

Ptychoptera  is  a  small  genus  including  sixteen  described  species,  almost 
all  occurring  in  the  Holarctic  region,  with  a  few  species  in  India  and 
Formosa.  Two  species  are  found  in  tropical  Africa,  and  another,  an 
undescribed  form,  in  southern  South  America;  hence  the  genus  is  probably 
found  thruout  the  world  in  countries  in  the  temperate  zones,  with  the 
probable  exception  of  Australia. 

The  literature  on  the  immature  stages  of  this  genus  is  summarized 
under  the  family  account. 

Topsent  (1914-16)  has  furnished  the  best  account  of  any  species  of 
the  genus  Ptychoptera.  His  observations  refer  to  Ptychoptera  albimana. 
The  following  notes  are  extracted  from  Topsent's  paper: 

The  eggs  range  in  number  from  520  to  587,  averaging  554.  They  are  pale  yellow,  slightly 
arcuated,  the  surface  curiously  ornamented,  the  dimensions  being  0.825  by  0.264  millimeters. 
The  duration  of  the  egg  stage  is  seven  days.  The  newly  hatched  larva  measures  3.85  milli- 
meters, the  respiratory  tube  0.99  millimeters.  The  adult  larva  measures  77  millimeters, 
of  which  the  respiratory  tube  is  20  millimeters.  The  growth  of  the  larva  is  rapid.  When 
fifteen  days  old  it  measures,  when  extended,  25  millimeters.  When  sixty-seven  days  old, 
it  measures  45  millimeters.  The  pupal  stage  is  from  ten  to  twelve  days,  averaging  eleven 
days.  There  are  normally  two  generations  a  year. 

De  Meijere  (1916:188-191)  gives  a  critical  account  of  the  structure 
of  the  larva  of  a  European  Ptychoptera.  It  agrees  well  with  the  description 
of  P.  rufodncta  given  below,  the  most  conspicuous  differences  shown  by 
De  Meijere's  species  being  as  follows:  labium  with  the  submentum  having 
the  sides  straighter;  mentum  with  the  anterior  margin  evenly,  but  gently, 
convex,  with  only  eighteen  evident  teeth.  The  details  of  the  maxilla, 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  775 

especially  of  the  setae  on  the  cardines,  are  not  the  same.  However, 
the  generic  characters  given  above  include  all  the  species  to  which  the 
writer  has  been  able  to  refer. 

Malloch  (1915-17  b:  240-241)  describes  and  figures  an  American  Ptychop- 
tera  which  is  presumably  P.  lenis  O.  S.  It  agrees  closely  with  the  other 
forms  in  most  respects.  The  author  indicates  the  mentum  as  having 
three  distinct  parts,  the  median  part  being  projected  beyond  the  level  of 
the  lateral  parts  and  having  more  teeth  (eight  instead  of  six).  Malloch's 
specimens  were  taken  in  the  Yellowstone  Park,  Wyoming,  in  August, 
1890.  From  the  locality  data  it  is  almost  certain  that  the  larvae  belong 
to  the  common  Ptychoptera  lenis  of  the  western  United  States. 

Tonnoir  (1919)  describes  a  curious  sexual  dimorphism  in  the  genus 
Ptychoptera. 

Ptychoptera  rufocincta  0.  S. 

1859     Ptychoptera  rufocincta  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  252. 

The  species  Ptychoptera  rufocmcta  is  common  around  wet  swales 
and  among  open  shrubbery.  The  adult  flies  are  on  the  wing  from 
May  to  early  July,  and  again  .in  late  August  and  early  September;  they 
may  be  double-brooded.  In  appearance  they  are  conspicuously  unlike 
their  relatives  of  the  Bittacomorphinae,  rather  resembling  certain  large 
fungus  gnats  (Mycetophilidae) . 

The  larvae  live  in  situations  similar  to  those  frequented  by  Bittaco- 
morpha,  and  often  occur  in  the  same  associations  —  with  larvae  of 
Limnophila  (Lasiomastix)  macrocera,  Pilaria  tenuipes,  P.  recondita,  and 
Erioptera  chlorophylla,  as  well  as  with  many  Chironomidae,  leeches, 
nematode  worms,  and  mollusks.  In  appearance  they  are  strikingly 
different  from  the  larvae  of  Bittacomorpha,  being  pale,  white  or  yellowish 
white,  with  the  body  almost  smooth,  not  covered  with  the  dense  trans- 
verse rows  of  setiferous  tubercles  found  in  Bittacomorpha,  and  with  the 
pseudopods  on  the  basal  abdominal  segments  poorly  developed.  The 
larvae  of  this  species  are  smaller  than  those  of  Bittacomorpha,  with  the 
head  proportionately  much  smaller.  In  structural  details,  however, 
they  are  rather  similar  to  the  larvae  of  Bittacomorpha.  The  larvae  of 
Ptychoptera  rufocincta  feed  on  the  decaying  vegetable  matter  in  their 
haunts.  When  fully  grown,  the  pupa  forms  within  the  last  larval  skin, 
and  the  elongate  pupal  spiracle  is  coiled  about  the  mesothorax,  as  described 

• 


77G  CHAELES  PAUL  ALEXANDER 

for  Bittacomorpha  clavipes  (Hart,  1898  [1895] :  19  L).  Careful  breedings  of 
this  species  in  1913  at  Orono,  Maine,  placed  the  indoor  pupal  life  at  four 
days  and  eighteen  hours,  and  that  in  nature  at  probably  five  days  —  an 
unusually  short  pupal  duration. 

Larva.—  Length,  when  fully  grown,  30-32  mm.;  when  fully  extended,  about  35  mm. 
Diameter,  2-2.2  mm. 

Head  light  reddish  brown,  not  marked  with  darker  spots  as  in  Bittacomorpha;  body 
whitish  or  pale  yellow;  tomentum  short,  pale;  seventh  and  eighth  segments  of  abdomen, 
and  breathing  tube,  light  brown. 

Body  almost  smooth,  sparsely  clothed  with  short,  appressed  hairs  arranged  in  indistinct 
transverse  rows,  on  intermediate  segments  of  body  there  being  about  twenty  of  these  rows; 
body  not  at  all  tuberculate,  as  in  Bittacomorpha.  Prothoracic  segment  short,  not  so  long 
as  mesothorax;  metathorax  nearly  as  long  as  preceding  two  segments  combined.  (In  older 
larvae  that  are  about  to  pupate,  the  right  pupal  breathing  horn  may  be  seen  coiled  under- 
neath the  skin  of  the  mesothorax.)  First  five  abdominal  segments  swollen  posteriorly  into 
a  ring  that  completely  surrounds  the  segments;  first  three  abdominal  segments  with  low, 
indistinct  pseudopods  on  either  side  of  the  median  line,  each  with  a  small,  recurved  claw; 
pseudopods  after  first  pair  more  widely  separated  than,  and  not  so  well  developed  as,  in 
the  Bittacomorphinae.  Abdominal  segments  4  and  5  more  elongate,  swollen  posteriorly 
but  not  bearing  pseudopods;  segment  6  narrowed  behind,  with  a  few  scattered,  outspreading 
hairs;  segment  7  narrower  than  preceding  and  telescopic  within  itself,  a,t  about  two-fifths 
the  length  there  being  a  transverse  row  of  long  hairs  marking  the  limit  of  telescoping;  when 
fully  extended,  segment  7  a  little  longer  than  segment  8;  segment  8  a  little  narrower,  and 
telescopic  basally  into  segment  7.  (The  parts  of  segments  7  and  8  which  are  exposed  in 
the  retracted  condition  are  brown  and  subchitinized,  and  bear  scattered,  outspread  hairs 
which  are  most  numerous  near  the  caudal  end  of  the  exposed  part  and  here  form  transverse 
rows;  similar  rows  of  sparse,  setiferous  punctures  are  on  the  dorsum  of  the  swellings  on  the 
first  five  abdominal  segments.)  Breathing  tube  (segment  9)  retractile  into  segment  8  for 
about  one-fourth  its  length;  when  retracted,  completely  concealing  gills.  Tracheal  gills 
two,  elongate-cylindrical,  situated  near  base  of  segment  9;  in  normal  position  of  rest,  gills 
usually  projecting  about  one-half  their  length  beyond  end  of  segment  8.  Apex  of  breathing 
tube  truncated  but  without  well-defined  lobes. 

Head  small,  proportionately  much  smaller  than  in  Bittacomorpha;  oval  to  somewhat 
pear-shaped;  narrow  anteriorly,  broadened  behind,  near  posterior  margin  abruptly  narrowed. 
Prefrons  broad,  conspicuous.  (As  noted  by  most  earlier  writers  on  the  genus  Ptychoptera,  the 
head  bears  numerous  punctures  with  setae  which  are  plumose  or  havo  a  branched  appearance 
[Plate  XV,  20].  The  writer  has  examined  numerous  specimens  under  high  magnification 
and  is  inclined  to  believe  rather  that  in  some  cases  several  bristles  arise  from  a  single  puncture 
and  are  closely  approximated  basally,  altho  free  distally,  and  that  the  plumose  appearance 
is  here  merely  apparent.  In  other  cases,  however,  actual  basal  fusion  has  taken  place.  The 
number  of  free  tips  from  a  puncture  varies  from  three  to  six,  five  and  six  being  common 
numbers.)  Labrum  broad,  transverse,  on  disk  two  large  setiferous  punctures  bearing 
branched  setae;  on  either  side  beneath,  conspicuous  tufts  of  long  hairs,  these  tufts  continued 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  777 

obliquely  proximad  to  near  median  line,  where  there  is  a  median  lobe  densely  covered  with 
long  hairs.  Epipharynx  small,  subquadrate,  margin  almost  transverse,  with  about  seven 
blunt  teeth;  sides  of  organ  with  long  hairs  which  are  directed  backward.  Labium  (Plate 
XIV,  12)  with  submentum  much  narrowed  on  basal  half,  sides  subparallel,  thence  expanded 
so  that  cephalic  end  is  about  twice  as  wide  as  caudal  end;  anterior  lateral  margins  produced 
into  blunt  projections;  mentum  (Plate  XV,  17)  broadly  subquadrate,  anterior  margin  nearly 
transverse  and  with  from  twenty  to  twenty-two  teeth  forming  an  irregular  comb.  (There  is 
considerable  variation  in  the  shape  of  this  comb  and  in  the  form  of  the  individual  teeth  com- 
prising it;  usually  the  median  third  of  the  mentum  is  produced  outward  beyond  the  lateral 
parts,  but  this  condition  is  not  always  well-marked;  the  individual  teeth  may  be  blunt  or 
acute.  In  the  specimen  shown  the  normal  condition  is  illustrated;  the  central  lobe  includes 
six  teeth,  and  each  lateral  part  about  seven  teeth,  the  lateral  teeth  being  usually  a  little  the 
larger;  in  some  specimens  the  median  lobe  has  the  teeth  very  indistinct,  while  in  others 
there  is  a  conspicuous  tendency  for  the  median  pair  to  unite  into  a  single  broad  tooth.) 
Prementum  pale  basally,  indistinctly  covered  with  pale  papillae;  palpi  lying  close  together, 
subparallel,  the  lateral  parts  with  short,  dense  hairs,  the  ventral  face  with  shorter  papillae; 
each  lobe  bearing  at  tip  a  small,  blunt,  cylindrical  knob  which  is  slightly  chitinized.  Antenna 
(Plate  XV,  18)  short,  almost  cylindrical  but  slightly  narrowed  basally,  bearing  on  truncate 
apex  about  five  sensory  papillae  of  various  diameters,  one  being  much  the  largest,  two  others 
being  long  and  very  slender;  one  of  the  papillae  is  bisegmented,  the  apical  part  being  more 
slender  than  the  basal  part.  (De  Meijere  [1916,  fig.  14]  shows  his  European  Ptychoptera 
as  having  the  antennae  two-segmented  and  with  the  sensory  papillae  very  different  from 
the  condition  found  in  P.  rufocincfa.)  Mandible  (Plate  XV,  19)  strong  and  powerful;  cutting 
edge  sub  triangular,  with  three  large  outer  teeth  and  about  six  or  seven  small  inner  ones; 
of  the  larger  teeth  the  outermost  is  the  slenderest,  the  third  is  the  largest  and  stoutest;  small 
inner  teeth  subequal  in  length,  the  outermost  stout,  inwardly  the  teeth  becoming  more 
slender;  mandible  on  ventral  face  near  margin  with  two  powerful  setae,  the  posterior  one 
often  recurved,  the  anterior  one  directed  forward.  Maxilla  (Plate  XV,  19)  with  cardines 
roughly  elongate- triangular,  proximal  angle  acute,  ventral  face  with  three  large  setiferous 
punctures  bearing  several  bristles  of  unequal  length  (the  writer  has  not  been  able  to  locate 
setae  on  the  middle  puncture) ;  outer  edge  of  sclerite  with  a  fringe  of  long  hairs,  longest  at 
narrow  inner  end  of  segment;  stipites  roughly  triangular,  with  a  group  of  short  spines  at 
apex;  palpi  antenniform,  stout,  cylindrical,  with  sensory  papillae  at  tip  (one  large  papilla, 
about  three  of  medium  size,  and  three  or  four  small  ones) ;  outer  lobe  of  maxilla  with  a  small, 
egg-shaped  or  subcylindrical,  knob  at  its  outer  angle,  below  which  the  rounded  lobe  is 
densely  clothed  with  long,  pale  hairs;  caudad  of  these,  along  margin,  a  row  of  from  six  to 
eight  powerful  spines  and  a  few  long  hairs. 

Pupa.—  Total  length,  34.3-40  mm. 

Length  excluding  kreathing  tube,  14.5-15.8  mm. 
Length  of  breathing  tube  alone,  19.8-26  mm. 
Width  of  body,  d.-s.,  1.6-1.7  mm. 
Depth,  d.-v.,  1.7-1.8  mm. 

Pronotal  breathing  horns  reddish  brown,   dark  brown  at  extreme  base;  thorax,  wing 
sheaths,  and  leg  sheaths  dark  brown;  abdomen  whitish,  with  small  tubercles    and  broad 


778  CHARLES  PAUL  ALEXANDER 

chitinized  apices  to  segments  dark  brown.  Pupa  most  readily  distinguished  from  pupa 
of  Bittacomorpha  by  the  venation,  the  position  of  the  tarsal  sheaths,  and  the  short,  non- 
stellate  abdominal  tubercles.  (Plate  XIV,  13.) 

Anterior  cephalic  crest  small,  deeply  bilobed  by  a  broad  V-shaped  notch,  the  rounded 
lobules  roughened  and  each  terminated  by  a  single  long  hair.  Dorsad  of  these,  two  smaller 
and  slenderer,  very  widely  separated,  tubercles,  each  lying  just  inside  antennal  sheaths. 
Antennal  bases  located  on  ventral  side  of  head,  between  eyes;  antennae  bent  dorsad  and 
thence  caudad  around  knee  joints  of  fore  legs,  the  tips  lying  just  outside  middle  tibiae;  apical 
antennal  segments  showing  distinctly  thru  sheaths.  Sheaths  of  maxillary  palpi  elongate, 
tips  curved  over  knee  joints  of  fore  legs.  Clypeus  smaller  and  narrower  than  the  conspicuous 
labium,  transversely  wrinkled.  Labium  rectangular,  each  half  with  tips  obliquely  truncated. 
(Plate  XIV,  14.) 

Pronotum  with  lateral  ventral  angles  almost  square,  each  with  about  two  small  setae. 
Breathing  horns  very  unequal  in  length;  the  right  one  greatly  elongated,  much  longer  than 
remainder  of  body,  enlarged  at  extreme  base,  the  outer  part  with  numerous  papillae  which 
are  more  numerous  and  more  approximated  toward  end  of  organ,  these  papillae  lying  in  a 
single  straight  line  which  makes  a  long  spiral  around  organ;  left  breathing  horn  very  small 
and  degenerate,  only  a  little  longer  than  sheath  of  maxillary  palpi,  curved,  with  about  a 
dozen  papillae  which  are  closely  crowded  toward  apex  of  organ.  Mesonotum  finely  and 
transversely  wrinkled,  prescutum  with  a  very  acute  V-shaped  dorso-median  mark,  its  apex 
directed  backward;  scutal  lobes  projecting,  each  with  a  few  tiny  hairs.  Postnotum  with 
two  small  tubercles  at  about  two-thirds  its  length,  one  on  either  side  of  a  pale  median  line. 
Metanotum  short,  sheath  of  halteres  extending  just  beyond  base  of  second  abdominal 
segment.  Wing  sheaths  clearly  showing  venation,  the  branched  media  being  characteristic 
of  the  genus;  wing  sheaths  extending  almost  to  end  of  second  abdominal  segment.  Leg 
sheaths  with  tips  of  fore  and  middle  tibiae  enlarged,  and  with  inner  apical  angle  of  each 
produced  into  long  points  for  the  long  tibial  spurs  of  adult  flies.  Leg  sheaths  extending  to 
just  before  end  of  third  abdominal  segment;  all  six  legs  lying  side  by  side,  not  overlapping 
as  in  Bittacomorpha  (Plate  XV,  21). 

First  abdominal  segment  chitinized  above,  apical  half  with  a  few  weak  tubercles.  Dorsum 
of  segments  2  to  6  with  posterior  margins  each  having  a  chitinized  band  set  with  about 
twenty-five  to  thirty  setiferous  tubercles,  the  outermost  ones  being  the  largest;  remainder 
of  dorsum  of  each  segment  with  irregular  transverse  rows  of  scattered  tubercles,  on  narrow, 
interrupted,  chitinized  bands;  usually  one  of  the  bands,  at  about  midlength  of  segment, 
broader  and  more  strongly  chitinized  than  the  others;  these  bands  obliterated  on  posterior 
segments,  but  caudal  band  here  very  wide;  about  fifteen  of  these  rows  on  segments  2  and 
6,  and  from  twenty  to  twenty-five  rows  on  segments  3  to  5;  these  bands  not  regularly  trans- 
verse, but  anastomosing  rather  freely,  not  occupying  more  than  half  of  abdominal  surface; 
segment  7  with  the  broad  caudal  band  only;  tubercles  rather  short,  with  three  or  four  short, 
irregular  spines  at  tip,  these  not  presenting  a  stellate  appearance  as  in  Bittacomorpha. 
Abdominal  sternites  similar  to  dorsum,  tubercles  lacking  where  leg  sheaths  rest  against 
segments  2  and  3;  segments  4  to  6  with  transverse  rows  similar  to  those  of  dorsum  but 
weaker;  caudal  bands  on  segments  4  to  7  very  broad,  the  caudal  margin  with  tubercles, 
there  being  about  twenty  tubercles  on  segment  4,  the  number  gradually  decreasing  to  seg- 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  779 

ment  7,  on  which  there  are  about  fifteen.  Abdominal  pleurites  with  tubercles  arranged  in 
longitudinal  rows,  there  being  about  six  rows  on  each  of  segments  2  to  6,  the  rows  being 
almost  continuous  for  the  whole  length  of  abdomen;  tubercles  at  caudal  margin  of  each 
segment  enlarged  and  powerful;  on  segment  6  the  rows  converging  behind  into  a  single 
powerful  tubercle  on  caudal  margin  of  segment;  a  similar  enlarged  tubercle  on  caudal  margin 
of  seventh  segment;  segment  7  and  cauda  narrowed.  Male  cauda  (Plate  XV,  22)  with  a 
prominent  dorso-median  lobe  projecting  directly  away  from  the  body;  dorsal  sheaths  short, 
indistinctly  bilobed;  ventral  sheaths  very  elongate,  divergent  (Plate  XV,  23).  Female  cauda 
(Plate  XV,  24)  with  the  same  prominent  dorso-median  lobe;  acidotheca  of  ovipositor  long, 
straight,  beyond  its  midlength  a  blunt,  conical  tubercle  directed  laterad  and  slightly  caudad; 
sternum  (Plate  XIV,  15)  with  ventral  lobe  only  about  half  length  of  tergal  acidothecae,  caudal 
margin  with  three  lobules. 

Nepionotype. —  Orono,  Maine,  June  24,  1913. 

Neanotype. —  With  the  larval  type. 

Paratypes. —  Both  larvae  and  pupae,  June  24  to  July  5,  1913. 

SUBFAMILY  Bittacomorphinae 

The  following  keys  separate  the  genera  of  the  subfamily  Bittacomor- 

phinae : 

Larvae 

Size  small  (total  length  under  20  mm.);  coloration  black,  breathing  tube  light  yellow; 
breathing  tube  entirely  retractile;  body  covered  with  very  long  projections  which  are 
incased  in  a  black,  horny  substance;  mandibles  with  an  inner  comb  of  teeth. 

Bittacomorphella  Alex.  (p.  779) 

Size  larger  (total  length  over  40  mm.);  coloration  rusty  red;  body  tapering  gradually  to  the 
long,  slender,  partly  retractile,  breathing  tube;  body  covered  with  transverse  rows  of  shorter, 
stellate  tubercles;  mandibles  without  an  inner  comb  of  teeth. 

Bittacomorpha  Westw.  (p.  783) 
Pupae 

Size  small  (length,  excluding  breathing  horn,  under  12  mm.);  right  breathing  horn  small, 
degenerate;  abdominal  tubercles  weak,  tipped  with  several  strong  setae. 

Bittacomorphella  Alex.  (p.  779) 

Size  larger  (length,  excluding  breathing  horn,  over  14  mm.);  right  breathing  horn  elongate, 
filiform,  longer  than  the  body;  abdominal  tubercles  strong,  elongate,  crowned  by  a  circlet 
of  four  or  five  spines  and  tipped  with  a  setiferous  papilla. .  .Bittacomorpha  Westw.  (p.  783) 

Genus  Bittacomorphella  Alexander  (Gr.  diminutive  of  Bittacomorpha) 

1916     Bittacomorphella  Alex.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  545. 

Larva. —  Body  short,  covered  with  very  prominent  projections  which  are  longest  on  lateral 
and  caudal  parts  of  body.  Pseudopods  prominent,  with  very  large,  curved  claws.  Breathing 
tube  short,  entirely  retractile  within  body.  Head  subquadrate,  the  foramen  ventral  in 
position;  setae  of  head  unbranched.  Mandible  with  an  inner  comb  of  teeth.  Mentum 
bilobed,  cephalic  margin  untoothed.  Coloration  black;  breathing  tube  light  yellow. 

Pupa.—  Right  breathing  horn  very  short,  degenerate.  Fore  tarsi  overlying  middle  tarsi. 
Tubercles  on  abdomen  moderately  elongated,  not  crowned  by  a  circlet  of  spines  as  in  Bitta- 
comorpha, but  tipped  with  a  long  seta. 


780  CHARLES  PAUL  ALEXANDER 

The  genus  Bittacomorphella  contains  but  two  described  species,  the 
genotype,  B.  jonesi  (Johns.),  and  the  larger  B.  sackenii  (Roder)  from 
western  America,  the  immature  stages  of  which  are  wholly  unknown. 
There  is  no  published  literature  on  the  biology  of  this  group  of  crane-flies. 

Bittacomorphella  jonesi  (Johns.) 

1905    Bittacomorpha  jonesi  Johns.     Psyche,  vol.  12,  p.  75-76. 

Bittacomorphella  jonesi  is  a  curious  little  phantom  crane-fly,  not  uncom- 
mon in  cold  Canadian  woods  thruout  the  Northern  States,  where  it  is 
usually  found  near  running  water  or  springs  and  often  in  small,  dark 
ravines  or  along  shaded  runs.  The  adult  flies  sometimes  lurk  beneath 
low,  dark  bridges  and  culverts,  where  they  are  often  associated  with 
species  of  Dolichopeza  and  Oropeza.  An  account  of  the  habits  of  the 
adults  of  this  species  may  be  found  in  an  earlier  paper  by  the  writer 
(Alexander,  19 16  b:  545-546). 

The  larval  habitat  is  very  different  from  that  of  other  local  species 
of  the  family,  which,  as  a  rule,  prefer  open  swamps,  swales,  or  wet 
meadows.  The  larvae  of  this  species  live  in  rich  organic  mud  in  shaded 
woods.  They  were  first  found  on  the  Bool  hillside  at  Ithaca,  New  York, 
beneath  decaying  beech  leaves  in  wet  or  damp  mud  which  was  mixed 
with  old  beechnuts,  hulls,  acorns,  butternuts,  broken  decayed  twigs, 
and  similar  debris.  The  Bool  area  is  a  very  steep  hillside  with  a  general 
northern  exposure,  heavily  shaded  with  tall  forest  trees.  In  former  days 
it  extended  far  to  the  eastward  and  was  connected  with  Slim  Jim  Woods, 
near  the  second  bridge  in  Forest  Home.  The  cut  area  is  now  a  pasture, 
but  patches  of  skunk  cabbage  and  cat-tails  still  persist.  On  the  shaded 
hillside  the  skunk  cabbage  occupies  pockets  or  level  areas  where  the  soil 
is  largely  calcareous.  The  forest  cover  consists  of  beech,  hard  maple, 
basswood,  yellow  birch,  red  oak,  butternut,  elm,  a  few  large  alders,  a 
few  aspens,  and  on  the  surrounding  hillside  a  thick  stand  of  hemlock. 
The  shrubs  include  Ribes  floridum  L'Her.,  Cornus  alternifolia  Linn,  f.,  and 
similar  species.  The  herbage  at  this  season  is  of  the  dominant  skunk 
cabbage,  young  seedlings  of  Impatiens  hi  flora  Walt.,  Geum  rivale  Linn., 
and  Cardamine  Douglassii  (Torr.)  Britt.  In  places  there  are  thick  mats 
of  mosses,  Brachythecium  rutabulum  (Linn.)  B.  &  S.,  on  the  limy  soil,  and 
Amblystegium  on  decaying  prostrate  limbs. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  781 

The  curious  larvae  of  Bittacomorphella  were  here  found  associated 
with  the  following  crane-fly  larvae  and  pupae:  Dicranomyia  stulta, 
Limnophila  adusta,  L.  (Dicranophragma)  fuscovaria,  Ulomorpha  pilosella, 
Penthoptera,  Molophilus  hirtipennis,  Erioptera  megophthalma,  Ormosia 
innocens,  0.  nigripila,  Tipula  collaris,  T.  oropezoides,  T.  cayuga,  and 
others.  In  addition  numerous  other  natural  associates  were  found,  such 
as  the  larvae  of  a  carabid  beetle,  Nebria  sahlbergi  Fisch.,  sow  bugs,  mollusks, 
and  a  great  variety  of  other  organisms.  On  July  10,  1914,  adults  of 
Bittacomorphella  were  not  uncommon  in  the  same  association,  at  which 
time  they  were  flying  with  other  crane-flies  such  as  Dicranoptycha  germana, 
Molophilus  hirtipennis,  M.  pubipennis,  Erioptera  vespertina,  E.  venusta, 
and  Gonomyia  blanda. 

The  first  larvae  of  Bittacomorphella  were  found  on  May  11,  and  at 
that  time  were  almost  fully  grown.  They  present  a  very  remarkable 
appearance,  being  black  or  very  dark  in  color  and  covered  with  numerous 
long  projections.  The  breathing  tube,  which  is  capable  of  entire  retraction 
within  the  body,  is  very  short,  and  is  light  yellow,  in  contrast  with  the 
remainder  of  the  body.  The  larvae  are,  as  a  rule,  very  slow  and  sluggish 
in  their  movements,  but  when  disturbed  they  become  more  active.  Large 
and  small  larvae,  of  two  distinct  sizes  only,  were  often  found  in  the  same 
situations  at  the  same  time.  A* fully  grown  larva  was  placed  in  rearing 
on  June  11,  1917,  and  emerged  as  an  adult  male  on  June  24.  This  pro- 
vides for  a  pupal  duration  of  not  more  than  thirteen  days,  but  the  pupal 
stage  is  undoubtedly  much  shorter. 

Larva. —  Total  length,  15-17  mm. 

Length  exclusive  of  breathing  tube,  12.5-14  mm. 

Length  of  breathing  tube,  2.4-3  mm.;  to  base  of   gills,    1.3-1.8    mm.;    beyond 

gills,  1.1-1.2  mm. 
Greatest  diameter  across  body,  1.7-2  mm. 

Coloration  dark  brown  to  almost  black,  breathing  tube  light  yellowish;  in  young  indi- 
viduals and  occasional  older  specimens,  coloration  more  rusty. 

Body  short,  stout,  cylindrical  but  appearing  depressed,  covered  with  numerous  elongate 
projections.  Breathing  tube  very  short  and  capable  of  complete  retraction  within  body 
(Plate  XVI,  25).  Body  appearing  proportionately  broader,  and  tapering  more  abruptly  to 
breathing  tube,  than  in  other  species  of  the  family  herein  discussed,  this  appearance  being 
due  to  great  length  of  lateral  body  projections. 

The  most  notable  single  feature  of  larva  consisting  of  the  numerous  projections  from 
body  (Plate  XVII,  33),  these  being  incased  in  a  blackened,  horny  substance  which  is  some- 
what brittle.  Usual  shape  of  body  extensions  cylindrical,  slender,  and  generally  simple 


782  CHAELES  PAUL  ALEXANDER 

except  for  those  near  end  of  body,  which  are  asymmetrically  once-forked.  Basal  half  of  pro- 
jections heavily  chitinized,  but  distal  end  almost  transparent.  Entire  surface  of  projections 
beset  with  numerous  transverse  rows  of  short  hairs,  usually  about  nine  to  twelve  hairs  in  each 
row,  nine  and  ten  being  common,  these  hairs  doubtless  serving  to  hold  the  blackened,  horny 
covering  of  the  projection.  Sensory  papillae  (Plate  XVII,  34)  borne  at  or  near  apices  of  pro- 
jections, each  with  a  long  bristle;  these  papillae  narrowed  at  base,  thence  enlarged  to  form 
a  head,  on  which  bristle  is  inserted;  usually  one  or  two  bristles  to  each  projection,  but 
occasionally  an  additional  one  present,  which  is  much  smaller  and  degenerate.  Dorsal 
body  projections  occupying  transverse  rows  across  segments,  those  near  lateral  and  caudal 
parts  of  body  being  long  and  powerful,  those  on  median  region  being  short  and  degenerate; 
segments  of  thorax  and  abdomen  subdivided  into  false  segments,  these  transverse  rows 
occupying  caudal  margins  of  these  segments,  there  being  usually  from  four  to  six  of  the 
weak  projections  between  the  powerful  lateral  ones.  In  addition  to  these  projections,  sparse 
branched  hairs  lying  in  the  same  transverse  rows  (Plate  XVII,  35).  Projections  at  end  of 
body  surrounding  base  of  breathing  tube  all  long  and  powerful,  and,  as  stated  above,  some 
weakly  bifurcate.  Ventral  body  projections  similar  to  those  of  dorsumbut  relatively  smaller. 

Abdominal  pseudopods  feebly  chitinized  at  tips,  with  very  large,  slightly  curved  claws 
which  are  but  little  shorter  than  the  pseudopods  themselves  (Plate  XVII,  32).  Breathing 
tube  short  and  stout,  surface  before  apex  transversely  wrinkled.  Papillae  rather  numerous, 
bearing  sense  hairs  scattered  over  surface  of  tube,  those  just  back  of  apex  short  and  spine- 
like,  those  farther  back  long  and  slender,  very  like  and  homologous  to  the  bristles  terminating 
the  body  projections,  as  discussed  above.  Gills  two,  stout,  about  one-third  length  of  terminal 
section  of  breathing  tube.  Body  projections  incrusted  with  a  black,  horny  substance,  as 
discussed  above;  on  lateral  projections,  apical  setae  likewise  incased,  at  least  basally,  pro- 
ducing a  bilobed  or  even  a  trilobed  appearance;  this  black  corneous  incrustation  brittle  and 
easily  removed,  leaving  projection  and  bristles  intact. 

Head  subquadrate,  sides  nearly  parallel,  occipital  foramen  ventral  in  position  (Plate  XVI, 
26);  surface  of  head  covered  with  numerous  small,  chitinized  points  which  are  longest  on 
posterior  angles;  anterior  ventral  angle  slightly  produced;  two  setiferous  punctures  on 
ventral  surface,  one  on  either  side  just  behind  anterior  angles,  the  other  closer  to  foramen. 
Dorsum  with  setae  arranged  as  shown  in  Plate  XVI,  27;  setae  simple.  Labrum  (Plate  XVII, 
31)  with  four  dorsal  setiferous  punctures  along  anterior  margin,  the  median  pair  closely 
approximated;  another  powerful  seta  occupying  each  lateral  angle  on  dorsal  side;  dense 
brushes  of  long  hairs  on  either  side  beneath.  Epipharynx  (Plate  XVII,  31)  supported  by  two 
powerful  chitinized  arms  connecting  across  midventral  region  and  then  extending  laterad 
and  expanding  outwardly  to  form  posterior  margin  of  labrum;  surface  of  epipharynx  with 
closely  appressed  teeth.  Labium  (Plate  XVI,  28)  with  cephalic  margin  of  mentum  almost 
entire,  the  broad  median  part  produced  cephalad  and  feebly  bilobed;  palpi  with  a  dense 
fringe  of  long  hairs  around  base  and  with  sensory  papillae  at  tips.  Antenna  (Plate  XVI, 
30)  short,  cylindrical,  somewhat  globular  or  barrel-shaped;  about  four  elongate  papillae 
and  two  or  three  shorter  ones,  these  papillae  terminal  in  position,  the  largest  one  occupy- 
ing the  inner  side.  Mandible  (Plate  XVI,  29,  30)  with  the  outer  angle  a  powerful  hook 
bearing  smaller  teeth  on  ventral  face  at  about  midlength;  inner  angle  flattened,  and,  besides 
terminal  blade,  bearing  a  comb  of  about  five  teeth,  the  innermost  being  the  longest  and 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  783 

slenderest;  a  dorsal  line  of  strong  setae  extending  from  base  of  outer  hook  inward;  outer 
edge  of  mandible  with  two  strong  setae,  these  protected  at  their  bases  by  small  dorsal  ears, 
or  projections,  from  the  mandible;  dorsal  face  of  mandible  with  a  powerful  hinged  prostheca; 
near  base  of  mandible  on  dorsal  face  a  curious  five-lobed  sensory  organ.  Maxilla  (Plate 
XVI,  29)  with  the  cardo  triangular,  the  ventral  or  outer  face  with  two  closely  approximated 
seti Serous  punctures;  stipes  triangular,  with  three  strong  setae  near  palpus;  palpus 
antenniform,  cylindrical,  rather  elongated,  and  with  about  six  sensory  papillae  at  tip, 
one  being  much  longer-than  the  others;  outer  lobe  of  maxilla  with  cephalic  margin  blackened 
and  chitinized,  inner  angle  with  a  dense  brush  of  long  hairs. 

Pupa. —  Length  excluding  breathing  horn,  9.5  mm. 
Width,  d.-s.,  1.7  mm. 
Depth,  d.-v.,  1.4  mm. 

The  following  description  is  taken  from  the  cast  pupal  skin  of  the  only 
specimen  that  was  reared: 

Antennal  bases  approximated  on  front  between  eyes.  Clypeus  bluntly  rounded  at  apex, 
transversely  wrinkled.  Lobes  of  labium  (Plate  XVII,  36)  broad,  rounded  apically;  maxillary 
palpi  short  and  stout,  broad  at  base,  gradually  narrowed  to  the  short  tip,  which  is  not 
recurved.  (The  structure  of  the  head  and  the  eyes  indicates  some  peculiar  characters  not 
possessed  by  the  pupae  of  related  genera,  but  the  cast  pupal  skin  is  insufficient  for  accurate 
diagnosis.)  Two  bristles  below  eye  and  just  above  base  of  palpus,  and  a  longer  and  more 
slender  seta  farther  laterad.  Sides  of  head  behind  antennae  appear  to  be  produced  laterad 
into  blunt  points.  Right  breathing  horn  small,  degenerate,  much  curved.  (In  the  single 
pupal  skin  available,  it  cannot  be  determined  whether  the  left  horn  has  been  broken  off  or  is 
undeveloped.)  Just  laterad  of  each  breathing  horn  a  small  tubercle  bearing  a  long  seta. 
Scutal  lobes  with  about  four  stout  setae.  Tarsal  sheaths  of  fore  legs,  as  in  the  subfamily, 
overlying  the  middle  pair  but  shorter  (Plate  XVII,  37). 

Abdomen  with  chitinized  bands  extensive,  as  in  Bittacomorpha.  Arrangement  of  tubercles 
about  as  in  other  species  of  family.  Tubercles  shorter  and  weaker  than  in  Bittacomorpha 
and  not  crowned  by  a  circlet  of  spines,  each  being  tipped  with  one  or  more  (four  or  five)  long 
setae  (Plate  XVII,  38);  the  pleura!  tubercles  the  longer  and  many  of  them  multisetose. 
Cauda  of  male,  in  general  features,  similar  to  that  in  Bittacomorpha,  the  dorsal  median 
lobe  (Plate  XVII,  39)  stout,  the  ventral  horns  (Plate  XVII,  40)  short  and  powerful, 
directed  laterad;  horns  on  dorsal  lobes  apparently  lacking. 

Nepionotype. —  Ithaca,  New  York,  May  30,  1917. 
Neanotype. —  Cast  pupal  skin,  reared  June  23,  1917. 
Paratypes  —  Topotypic,  May  15  to  June  10,  1917. 

Genus  Bittacomorpha  Westwood  (Gr.  Bittacus  +  shape) 

1835    Bittacomorpha  Westwood.     London  and  Edinburgh  Phil.  Mag.,  vol.  6,  p.  281. 

Larva. —  Form  elongate,  body  gradually  narrowed  behind  into  the  partly  retractile 
breathing  tube.  Integument  with  transverse  rows  of  tubercles.  Pseudopods  on  abdominal 
segments  1  to  3  prominent,  with  large  curved  claws.  Head  subpyriform,  dorsum  with  rows 


784  CHARLES  PAUL  ALEXANDER 

of  prominent  black  spots  converging  behind.     Mandible  stout,  with  a  single  powerful  outer 
tooth.     Mentum  bilobed,  anterior  margin  not  comblike.     Color  of  body,  rusty  red. 

pupa,~  Right  pronotal  breathing  horn  very  elongate;  the  left  very  small  and  short, 
subdegenerate.  Fore  tarsi  overlying  middle  tarsi.  Tubercles  on  abdomen  very  long,  located 
on  broad  transverse  bands  of  chitin,  each  tubercle  with  a  star  of  four  or  five  spines  surrounding 
the  apex,  which  bears  a  long  seta. 

The  genus  Bittacomorpha,  as  here  restricted,  includes  but  two  species  - 
the  genotype,  B.  clavipes   (Fabr.),  and   B.  occidentalis    Aid.  of  western 
America,   concerning  the  biology  of   which  nothing  has    been   recorded. 
The  literature  on  the  immature  stages  of  Bittacomorpha  clavipes  is  sum- 
marized under  the  family  account  (page  773). 

Bit'acomorpha  clavipes  (Fabr.) 

1781     Tipula  clavipes  Fabr.     Spec.  Ins.,  vol.  2,  p.  404. 

1835    Bittacomorpha  clavipes  Westw.   London  and  Edinburgh  Phil.  Mag.,  vol.  6,  p.  281. 

Bittacomorpha  clavipes,  the  "  phantom  crane-fly,"  is  a  common  and 
widely  distributed  species  thruout  North  America  east  of  the  Rockies. 
It  is  easily  recognized  by  the  black-and-white-banded  legs,  with  their 
conspicuously  enlarged  and  swollen  metatarsi.  The  species  is  very 
characteristic  of  alder  swamps  and  the  wet  margins  of  ponds.  While  in 
copulation  the  insects  often  fly,  the  female  ahead,  the  male  trailing  on 
behind  like  the  tail  of  a  kite.  When  they  alight  on  a  plant  stem,  the 
female  is  invariably  uppermost,  the  male  often  hanging  free  with  none 
of  its  feet  on  a  support.  The  swollen  metatarsi  are  almost  completely 
filled  by  the  tracheae,  and  these  serve  to  buoy  the  insects  as  they  drift 
about  in  the  wind.  Brues  (1900)  describes  these  peculiar  tracheal  dilations 
in  detail.  He  says,  in  part: 

When  flying,  Bittacomorpha  uses  the  wings  scarcely  at  all,  relying  in  great  measure  upon 
wind  currents  for  transportation.  The  legs  are  exceedingly  light,  as  the  exoskeleton  is  thin 
and  delicate,  and  encloses  practically  no  tissue  which  can  serve  to  increase  their  weight. 

In  a  letter  from  Dr.  J.  G.  Needham,  dated  September  27,  1917,  valuable 
data  on  this  habit  of  drifting  are  furnished,  as  follows: 

Yesterday  while  crossing  the  Fall  Creek  bridge  near  my  home  on  Cornell  Heights,  I  made  an 
observation  on  Bittacomorpha  that  interested  me  greatly.  A  breeze  was  blowing  up  the  gorge, 
and  on  the  breeze  a  Bittacomorpha  was  drifting  rapidly  upward  in  the  usual  flight  attitude, 
with  broadly  outspread  legs,  the  swollen  metatarsi  hanging  vertically,  all  phantom-like  in 
slenderness  and  in  strongly  contrasting  black  and  white.  It  came  up  from  below  the  level 
of  the  rail,  swept  past  within  two  feet  of  my  face,  and  passed  on  upward  with  the  breeze  until 
lost  to  view,  perhaps  100  feet  higher  than  the  bridge,  and  much  farther  upstream.  Since 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  785 

the  creature  can  fly  only  very  slowly  and  here  was  moving  several  times  faster  (I  could  not 
see  whether  it  was  using  its  wings),  it  was  obviously  drifting  in  the  wind.  Perhaps  this  is  a 
normal  function  of  the  expanded  metatarsi. 

The  larvae  are  usually  abundant  in  decaying  vegetable  matter  in  rich 
organic  mud  about  ponds  and  in  swamps.  The  writer  has  found  them 
especially  numerous  in  the  Basin  Swamp  at  Orono,  Maine  (in  1913), 
and  near  Round  Pond  at  McLean,  New  York.  At  Orono  they  were 
associated  with  larvae  and  pupae  of  Ptychoptera  rufocinda,  Limnophila 
macrocera,  Pilaria  tenuipes,  P.  recondita,  Erioptera  chlorophylla,  a  variety 
of  chironomid  larvae,  numerous  larvae  of  Trichoptera  in  their  cases, 
nematodes,  and  leeches.  Needham  and  Betten  (1901:574-575)  give  a 
summary  of  the  larval  habitat  of  this  species  as  they  found  it  in  the 
northern  Adirondacks.  Weston  and  Turner  (1917:53)  have  recorded 
the  larvae  as  being  scavengers  and  thus  serving  as  important  factors 
in  the  elimination  of  sewage  in  the  Coweeset  Stream  near  Brockton, 
Massachusetts. 

The  immature  stages  of  this  interesting  crane-fly  have  been  well  con- 
sidered by  Hart  (1898  [1895]:  189-195),  whose  account  has  been  briefly 
summarized  by  Howard  (1912:95-96).  The  larvae  are  found  in  shallow 
water  that  is  filled  with  decaying  vegetable  matter.  Here  they  live  in 
the  mat  of  dead  stems  of  rushes,  grasses,  and  willow  leaves,  in  semi- 
stagnant  or  slowly  flowing  water.  The  larvae  are  elongate-cylindrical, 
with  a  long,  partly  retractile  breathing  tube.  They  are  deep  rusty  red 
or  brown  in  color,  quite  distinct  from  the  pale  whitish  larvae  of  Ptychoptera 
or  the  black  larvae  of  Bittacomorphella.  They  feed  on  decaying  vegetable 
matter,  diatoms,  and  mud  that  is  filled  with  organic  matter.  They  rest 
beneath  the  surface  of  the  water,  with  the  tip  of  the  extended  breathing 
tube  at  the  surface  film  or  just  beneath  the  surface,  in  the  latter  case 
breathing  by  means  of  the  small  tracheal  gills.  When  about  to  pupate, 
the  very  long,  coiled  breathing  tube  of  the  pupa  is  wound  around  the 
thorax  beneath  the  larval  skin.  On  pupation  the  tube  soon  straightens 
out  into  a  very  long,  stiff,  bristle-like  structure.  Like  the  larvae,  the 
pupae  rest  beneath  the  surface  of  the  water,  with  the  tip  of  the  breathing 
tube  projecting  above  the  surface  film.  The  pupal  duration  is  apparently 
about  a  week. 

Larva. —  Length  when  fully  extended,  about  60  mm. 
Length  of  breathing  tube,  about  20  mm. 
Diameter  of  body,  about  2.6  to  3  mm. 

4 


786  CHARLES  PAUL  ALEXANDER 

Body  tapering  gradually  at  either  end,  posterior  end  prolonged  into  breathing  tube. 
Usual  color  pale  rusty  brown,  but  the  writer  has  found  a  few  nearly  full-grown  specimens 
which  were  as  pale  in  color  as  the  larvae  of  Ptychoptera.  Body  covered  with  numerous 
transverse  rows  of  small  tubercles,  or  papillae,  which  bear  short  setae.  Head  broadly  ovate, 
convex  above,  where  it  is  conspicuously  marked  with  rows  of  black  spots,  these  interrupted 
lines  converging  behind.  Mouth  parts  in  general  similar  to  those  in  Bittacomorphella,  the 
main  points  of  difference  being  as  follows:  mandible  (Plate  XVIII,  42)  shorter  and  stouter, 
ending  in  a  powerful  outer  tooth,  the  comb  of  inner  teeth  being  reduced  to  about  eight  small 
tubercles,  the  two  bristles  on  outer  margin  of  mandible  not  overlapped  by  projecting"  ears  "; 
labium  (Plate  XVIII,  41)  shorter  and  stouter,  with  a  different  arrangement  of  papillae;  epiphar- 
vnx  long,  narrowed  behind,  distinctly  bilobed,  each  half  with  parallel  rows  of  long,  comblike 
teeth  projecting  proximad;  anterior  comb  of  epipharynx  with  the  anterior  teeth  the  largest, 
the  teeth  gradually  reduced  in  size  behind;  posterior  comb  with  the  rows  of  teeth  widely 
separated  anteriorly,  approximated  behind  so  as  to  be  contiguous  or  nearly  so  at  their  ends; 
space  between  these  rows  filled  with  long  hairs;  anterior  teeth  small  and  feebly  chitinized, 
posterior  teeth  stronger.  First  three  abdominal  segments  bearing  conspicuous  pseudopods, 
each  terminated  by  a  sharp,  slender  claw  which  fits  into  a  groove  on  the  face  of  the  pseudopod. 

Pupa.—  Total  length,  40-60  mm. 

Length  excluding  breathing  tube,  15.5-25  mm. 

Length  of  breathing  tube,  25-35  mm. 

Degenerate  breathing  tube,  length  2  mm.,  diameter  0.2  mm. 

Width,  d.-s.,  1.8  mm. 

Depth,  d.-v.,  2.6  mm. 

Breathing  tube  light  brown;  wing  sheaths  brown;  leg  sheaths  light  brownish  yellow  and 
dark  brownish  black,  alternated,  corresponding  to  the  leg  markings  of  the  adult  fly.  Abdomen 
pale  yellow,  rather  uniformly  covered  with  abundant  brownish  tubercles  and  transverse, 
chitinized  plates,  these  brown  areas  scarcer  on  pleura  and  not  especially  abundant  on  apical 
margins  of  segments. 

Pupa  somewhat  similar  in  general  structure  to  pupa  of  Ptychoptera.  Anterior  cephalic 
crest  small,  lobules  rounded,  each  tipped  with  a  long,  stout  seta;  immediately  behind  anterior 
crest,  a  similar  blunt,  bilobed  projection  of  front;  laterad  of  crest,  a  slender,  elongate  tubercle 
on  either  side,  immediately  behind  antenna!  sheaths,  each  with  a  long  seta;  two  other  setiferous 
tubercles  on  head  behind  antennae  and  maxillary  palpi.  Antennal  bases  approximated 
between  eyes.  Sides  of  head,  laterad  of  eyes,  with  a  small  setiferous  tubercle.  Maxillary 
palpi  not  recurved  at  tip,  as  in  Ptychoptera,  ending  opposite  knee  joint.  Clypeus  elongate, 
gradually  narrowed  toward  apex,  transversely  wrinkled;  two  hairs  toward  base  near  inner 
margin  of  eye.  Each  half  of  labium  broad,  roughly  subquadrate,  tips  broadened  and 
obliquely  truncated  (Plate  XVIII,  43). 

Breathing  horns  almost  as  in  Ptychoptera.  (Nearly  always  it  is  the  right  horn  that  is  elon- 
gated, but  in  about  ten  per  cent  of  the  specimens  the  left  horn  is  elongated  while  the  right 
is  degenerated;  Hart  records  one  specimen  in  which  both  horns  were  developed,  but  unequally, 
the  right  measuring  23  mm.  and  the  left  13  mm.;  some  of  the  specimens  recorded  by  Hart 
are  larger  than  any  that  the  writer  has  ever  seen.)  Wing  sheaths  ending  almost  opposite  tips 
of  fore  tarsi;  media  uubranched.  Leg  sheaths  (Plate  XVIII,  47)  with  fore  tarsi  much  shorter 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  787 

than  the  others  and  lying  directly  over  middle  tarsi;  tarsi  of  hind  and  middle  legs  parallel 
and  extending  beyond  tips  of  fore  tarsi.  Scutal  lobes  each  with  about  four  setiferous  tubercles. 
Thorax  and  first  abdominal  segment  transversely  crenulated. 

Abdomen  with  transverse  bands  of  chitin  much  broader  than  in  Ptychoptera,  so  that  they 
cover  almost  the  entire  abdominal  surface;  these  bands  with  about  twelve  tubercles  on 
segments  3  and  4,  about  ten  on  segment  5,  and  from  six  to  eight  on  the  posterior  segments; 
tubercles  of  various  sizes,  small  and  somewhat  degenerate  ones  being  interspersed  with 
larger  ones;  tubercles  long  and  slender,  each  crowned  by  a  circlet  or  star  of  from  three  to 
six  (usually  four  or  five)  stout  spines  (Plate  XVIII,  44-46),  a  setiferous  papilla  arising 
from  the  center  of  this  circlet  of  spines;  spines  on  pleura  longer  than  those  on  remainder  of 
abdomen,  but  not  arranged  in  distinct  longitudinal  rows  as  in  Ptychoptera,  being  usually  more 
irregular,  in  some  cases  showing  three  or  four  more  or  less  distinct  rows;  these  pleural  chitin- 
ized  areas  usually  bearing  from  two  to  four  tubercles,  which  are  closely  approximated  basally 
so  as  to  present  a  somewhat  branched  appearance.  Male  cauda  (Plate  XVIII,  48)  as  in 
Ptychoptera,  but  dorsal  median  lobe  very  short  and  stout;  tubercles  on  segment  immediately 
before  cauda  long  and  slender,  similar  to  those  on  remainder  of  abdomen. 

Nepionotype. —  Ithaca,  New  York,  May  15,  1917. 
Neanotype. —  Orono,  Maine,  June  24,  1913. 
Paratypes. —  With  the  type  pupa. 

Malloch's  figure  of  the  pupa  (1915-17  b:  pi.  35,  fig.  6)  is  diagrammatic. 
It  was  probably  made  from  a  female  individual,  the  antennal  sheaths 
being  shorter  in  this  sex  than  in  the  male. 

FAMILY  Rhyphidae 

Larva. —  Body  eucjphalous,  amphipneustic.  Mandibles  opposed.  Eyespots  distinct. 
Spiracles  on  sides  of  prothorax.  Thoracic  and  abdominal  segments  divided  by  false 
constrictions.  Spiracular  disk  surrounded  by  two  or  five  lobes '  (Rhyphinae)  or  by  four 
lobes  (Trichocerinae) ,  or  unprovided  with  lobes  (Mycetobiinae) . 

Pupa. — Head  with  a  bilobed  setiferous  cephalic  crest.  Palpi  stout,  straight.  Prono- 
tal  breathing  horns  short,  not  prominent.  Tarsal  sheaths  lying  in  pairs,  one  above 
another,  the  fore  legs  lying  on  the  middle  legs,  and  these  latter  on  the  hind  legs.  Lateral 
abdominal  spiracles  small  but  distinct. 

The  family  Rhyphidae  includes  an  apparently  heterogeneous  group 
of  genera  which  are  in  reality  very  closely  related.  The  adults  are  of 
diverse  appearance,  but  the  immature  stages  are  exceedingly  similar  to 
one  another  and  undoubtedly  all  three  of  the  groups  included  in  the 
family  are  closely  allied. 

The  immature  stages  of  the  Rhyphinae  (Rhyphus)  have  been  discussed 
by  many  entomologists,  among  others  by  Johannsen  (1910:35-36, 


788  CHARLES  PAUL  ALEXANDER 

Rhyphus  pundatus)  and  by  Malloch  (1915-17  b:  243,  R.  punctatus).  The 
larvae  are  often  handsomely  banded  and  mottled  with  brown  or  purplish. 
Johannsen  and  other  authors  describe  the  cauda  as  ending  in  two  short 
lobes,  but  Malloch  mentions  five  such  lobes.  The  general  structural 
characters  are  those  described  above  for  the  family.  The  larvae  occur 
in  decaying  vegetable  matter,  in  manure  (especially  horse  and  cow  dung), 
in  sewage,  and  in  similar  material. 

The  Mycetobiinae  are  represented  by  Mycetobia,  a  curious  fly  which 
superficially  resembles  a  mycetophilid  rather  than  a  crane-fly.  Long  ago 
Lyonet,  Dufour,  Guerin-Me"neville  and  others  described  and  figured  the 
larva  of  Mycetobia  and  noted  the  eucephalous  condition  of  the  head  and 
the  amphipneustic  spiracles.  Osten  Sacken  (1863)  first  suspected  the 
affinities  of  this  genus  with  Rhyphus.  More  recently,  work  by  Johannsen 
(1910:31-32),  Malloch  (1915-17  a,  and  1915-17 b: 244-245),  Edwards 
(1916),  Knab  (1916),  and  others  has  definitely  settled  the  relationship 
of  this  insect  with  the  Rhyphidae.  The  larvae  and  the  pupae  agree 
closely  with  the  general  family  characters  discussed  above.  The  larvae 
occur  in  decaying  wood  and  about  fermenting  sap  in  wounds  of  trees. 
The  genera  Ditomyia  Winn.  and  Symmerus  Walk,  are  now  placed  in  a 
separate  family  from  Mycetobia,  the  Ditomyiidae  (Keilin,  1919). 

Until  recently,  the  Trichocerinae  have  been  considered  as  being  mem- 
bers of  the  family  Tipulidae.  They  include  only  the  genus  Trichocera, 
with  about  twenty-five  nominal  species,  and,  presumably,  Ischnothrix 
Bigot,  represented  by  a  single  species  from  Cape  Horn.  From  the  general 
appearance  of  the  adult,  these  flies  have  usually  been  referred  to  the  tribe 
Limnophilini,  in  a  position  near  the  genus  Limnophila.  Brunetti  (1912) 
referred  them  to  the  Pediciini,  and  most  other  recent  workers  have  accorded 
them  tribal  or  subfamily  rank  in  the  Tipulidae.  Bezzi  (1914:214), 
influenced  by  the  work  of  Keilin  (1912),  referred  Trichocera  to  the 
Rhyphidae,  but  later  (1918a:20)  placed  it  back  in  the  Tipulidae  (as  Lim- 
nobiidae).  Malloch  (1915-17b:234)  likewise  places  Trichocera  with  the 
Tipulidae,  but  mentions  the  close  resemblance  of  the  larva  to  that  of 
Rhyphus.  The  best  discussions  of  the  morphology  of  the  larva  and  the 
pupa  are  those  by  Keilin  (1912)  and  De  Meijere  (1916:191-194),  both 
of  whom  were  strongly  impressed  by  the  striking  resemblance  of  the  larva 
to  that  of  Rhyphus.  In  the  present  paper,  the  Trichocerinae  is  the  only 
group  considered  in  detail. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  789 

SUBFAMILY  Trichocerinae 

Genus  Trichocera  Meigen  (Gr.  hair  +  horn) 

1800     Petaurista  Meig.     Nouv.  Class.  Mouch.,  p.  15  (nomen  nudum) 
1803     Trichocera  Meig.     Illiger's  Mag.,  vol.  2,  p.  262. 
1911     Paracladura  Brun.     Rec.  Indian  Mus.,  vol.  6,  p.  286. 

Larva.—  Body  eucephalous,  amphipneustic.  Thoracic  segments  divided  into  two  annuli. 
Spiracles  on  lateral  margin  of  posterior  ring  of  prothorax.  Abdominal  segments  divided 
into  three  annuli.  Cauda  ending  in  four  lobes,  ventral  lobes  the  longer  and  more  slender; 
lobes  bearing  numerous  stout  hairs  near  tips  on  outer  face.  Eyespots  distinct.  Lateral 
plates  of  head  widely  separated  on  midventral  line.  Mandible  with  prostheca  distinct. 

Pupa.—  Cephalic  crest  small,  lobes  with  stout  setae.  Clypeus  short;  labrum  dumb- 
bell-shaped; palpal  sheaths  stout.  Antenna  elongate.  Leg  sheaths  lying  in  pairs  above  one 
another,  gradually  lengthening,  fore  pair  the  shortest,  posterior  pair  the  longest.  Pronotal 
breathing  horns  short.  Abdominal  spiracles  small,  but  distinct  and  functional. 

The  small  winter  gnats  of  the  genus  Trichocera  are  rather  familiar, 
since  they  are  not  rare  during  the  winter  months  in  cellars  or  even  in 
the  open  on  warm  days,  occurring  in  sunlit  places  in  small,  dancing 
swarms.  They  are  abundant  during  fall  and  spring.  They  occur  also 
in  cool,  shady  places  in  summer,  but  are  less  in  evidence  at  this  season. 

Trichocera  is  found  somewhat  commonly  and  regularly  in  mines,  often 
at  very  considerable  depths.  Boheman  (1850)  records  specimens  of  T. 
regelationis  in  mines  600  feet  below  the  surface,  and  Lampa  (1890)  also 
records  the  species  as  being  found  at  considerable  depths.  Dr.  H.  B. 
Hungerford  found  numerous-  adults  of  a  species  of  Trichocera  in  the 
Amethyst  silver  mine  near  Creede,  Colorado,  in  1914.  Specimens  that 
he  obtained  were  taken  at  the  sixth  level,  but  the  miners  said  the  insects 
were  to  be  seen  in  all  parts  of  the  mine;  along  the  laterals  at  the  sixth 
level  they  were  noted  7000  feet  from  the  entrance.  It  is  supposed  that 
these  individuals  breed  in  the  animal  waste  which  naturally  accumulates 
in  such  places.  Trichocera  is  also  a  characteristic  inhabitant  of  caverns 
and  grottoes,  all  stages  being  found  in  such  situations  (Schmitz,  1909:80; 
Bezzi,  1911-12:46-47,  49,  and  1914:214). 

The  swarming  and  mating  of  these  flies  is  well  known.  It  has  been 
ably  described  by  Ainslie  (1907),  and  is  here  discussed  only  in  general 
terms.  The  insects  swarm  commonly  in  the  autumn.  Sometimes  the 
swarms  include  but  comparatively  few  individuals,  but  at  other  times 
many  thousands  participate.  They  swarm  usually  from  five  to  twenty- 
five  feet  above  the  ground,  all  facing  in  the  same  direction,  that  is, 


790  CHARLES  PAUL  ALEXANDER 

toward  the  wind  or  breeze.  Mating  takes  place  in  the  air,  and  united 
pairs  then  fly  away  or  drop  to  the  ground  beneath.  The  swarms  are 
often  very  dense,  and  individuals  come  in  frequent  contact  with  one 
another.  When  the  breeze  shifts,  the  swarm  immediately  readjusts  its 
position  and  direction. 

The  immature  stages  of  Trichocera  are  spent  in  decaying  vegetable 
matter,  beneath  dead  or  decaying  leaves,  in  debris,  in  fungi,  and  in 
similar  situations.  Sometimes  the  larvae  and  pupae  are  rather  numerous 
in  stored  roots  and  tubers,  especially  potatoes,  in  which  cases  they  may 
assume  an  economic  importance  (Johannsen,  1910:34-35;  Carpenter, 
1912).  The  specimens  used  by  the  writer  for  study  are  part  of  Johannsen's 
material,  determined  as  T.  regelationis  from  Patten,  Maine.  The  tax- 
onomic  condition  of  the  group  is  such  that  no  specific  identification  of 
the  adult  flies  can  be  attempted  at  this  time.  The  immature  stages  of  the 
generalized  subgenus  of  Trichocera,  Diazosma  Bergroth,  are  unknown. 

The  most  important  literature  on  the  genus  Trichocera  is  as  follows: 

Trichocera  regelationis General Dufour,  1840: 161. 

Trichocera  regelationis Larva,  general Schmitz,  1909:80,  pi.  8,  fig.  3. 

Trichocera  regelationis Larva,  pupa Johannsen,  1910:34-35,  figs.  51- 
ST. 

Trichocera  regelationis Pupa De  Meijere,  1916: 194. 

Trichocera  hiemalis Larva,  general Curtis,  1846 b. 

Trichocera  hiemalis General Cameron,  1917: 63. 

Trichocera  fuscata General Carpenter,    1912.     (Damage.) 

Trichocera  sp Larva Bremi-Wolf,  1846: 175. 

Trichocera  sp Larva Ferris,  1847: 37,  pi.  1,  fig.  3. 

Trichocera  sp General Bezzi,  1911-12:46-47,  49. 

Trichocera  sp Larva De  Meijere,  1916: 191-194,  figs. 

21-23. 

Trichocera  sp Larva,  pupa Keilin,  1912.     (Morphology.) 

Trichocera  sp Larva Malloch,  1915-17  b:  234-235,  pi. 

26,  fig.  1;306. 

Trichocera  regelationis,  supposition. 

Larva, —  Length,  8-9.5  mm. 
Diameter,  1  mm. 

Coloration  pale  brown  in  preserved  material,  whitish  in  fresh  specimens. 

Body  rather  short,,  cylindrical  to  slightly  depressed  (Plate  XIX,  49).  Pseudopods  lacking. 
Head  complete,  non-retractile,  strongly  chitinized;  lateral  plates  of  head  widely  separated 
on  midventral  line,  connected  only  by  a  narrow  bridge  posteriorly;  chaetotaxy  as  shown 
(Plate  XIX,  52  and  53).  Mandibles  opposed,  of  three  parts,  principal  segments  bearing 
on  inner  side  near  base  an  apparently  movable  appendage  (prostheca)  which  has,  besides  the 
large  apical  tooth,  three  smaller  teeth.  Labrum  bluntly  rounded,  with  long  hairs.  Epi- 
pharynx  with  lateral  combs  of  about  six  blunt  teeth.  Antenna  two-segmented;  basal  segment 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  791 

very  short,  disk- shaped,  inserted  on  a  Targe  brown-margined  plate  which  is  part  of  the  head 
chitin;  second  segment  much  narrower,  egg-shaped;  in  addition  to  this  segment  there  are 
several  small  sensory  papillae  on  the  end  of  the  first  segment. 

Segments  of  body  divided  into  secondary  annuli,  thoracic  segments  with  two  such  rings, 
abdominal  segments  with  three;  annuli  bearing  transverse  rows  of  short  setae.  Anterior 
spiracles  on  posterior  ring  of  prothorax  near  lateral  margin  conspicuous,  smaller  than  posterior 
spiracles  but  constructed  on  same  general  principle.  Spiracular  disk  surrounded  by  four 
lobes;  ventral  lobes  longer  and  more  slender  than  dorsal  pair,  inner  face  narrowly  chitinized, 
outer  face  densely  clothed  with  abundant  short  yellow  hairs  (Plate  XIX,  50  and  51); 
dorsal  lobes  shorter  and  blunter,  with  short  hairs  on  apices  of  outer  face.  Spiracles  large, 
at  base  of  dorsal  lobes. 

Pupa. —  Length,  7.5-7.8  mm. 
Width,  d.-s.,  1.4  mm. 
Depth,  d.-v.,  1.4  mm. 

Coloration  whitish;  head,  thorax,  and  sheaths  of  appendages  brown. 

Anterior  cephalic  crest  small,  lobes  widely  separated,  each  tipped  with  a  long,  stout  seta 
directed  ventrad;  a  tiny  seta  just  behind  each  anterior  lobe.  Antennal  bases  above  and 
slightly  between  the  eyes,  bent  dorsad  and  thence  caudad,  passing  behind  joints  of  legs,  in 
the  female  attaining  to  about  one-third  length  of  wing.  Frontal  region  between  eyes  slightly 
tumid,  somewhat  shiny.  Clypeus  short,  the  sides  parallel,  the  apex  U-shaped;  labrum 
broad,  dumb-bell-shaped,  the  caudal  margin  concave.  Maxillary  palpi  very  short  and  stout, 
ending  before  knes  joints  of  fore  legs  (Plate  XIX,  55).  A  small  tubercle  just  laterad  of 
base  of  antenna.  Each  cheek  produced  into  a  long,  blunt,  wrinkled  tubercle. 

Mesonotum  (Plate  XIX,  54)  strongly  gibbous,  pale  medially,  narrowed  in  front,  anterior 
margin  truncated  and  sending  a  sharp  median  carina  cephalad;  sides  of  mesonotum  opposite 
wing  root  with  four  small  setae,  in  two  slightly  separated  groups.  Anterior  angles  of  pro- 
notum  with  a  short  bristle.  Breathing  horns  small,  short  and  almost  straight,  broad  basally, 
apical  half  narrow,  inner  face  fused  or  closely  approximated  with  pronotum,  apex  cleft. 
Wing  sheaths  attaining  level  of  tips  of  fore  tarsi;  venation  rather  distinct.  Leg  sheaths  with 
fore  legs  stout;  fore  tarsi  overlying  middle  tarsi  (Plate  XIX,  55);  middle  tarsi  overlying 
hind  tarsi;  terminal  segments  of  tarsi  swollen.  Abdominal  segments  divided  into  about 
three  false  annuli;  caudal  margin  of  each  segment  fringed  with  short  hairs.  Tiny 
abdominal  spiracles  on  pleural  segments.  Female. ovipositor  (Plate  XIX,  56)  with  the  dorsal 
valves  short,  widely  separated,  acutely  pointed;  ventral  acidotheca  elongate,  approximated, 
bent  slightly  ventrad. 

Larvae  and  pupae. —  Patten,  Aroostook  County,  Maine,  May  3  and  23,  1907. 

t 

FAMILY  Tipulidae 

The  family  Tipulidae  is  the  largest  group  of  crane-flies,  and  possibly 
the  only  one  to  which  the  name  is  justly  applicable.  It  includes  a  vast 
number  of  species  (nearly  three  thousand),  arranged  in  about  one  hundred 


792  CHARLES  PAUL  ALEXANDER 

and  forty  genera.  The  species  are  found  in  most  parts  of  the  world,  being 
restricted  only  by  intense  heat  and  cold.  Crane-flies  require  moisture 
in  order  to  complete  their  development,  and,  as  a  consequence,  are  almost 
always  found  in  the  neighborhood  of  flowing  or  stagnant  water.  No 
species  known  to  the  writer  are  inhabitants  of  desert  conditions,  the 
nearest  approach  probably  being  some  Eriopterini,  such  as  Helobia, 
Trimicra,  and  other  genera. 

The  immature  stages  frequent  very  wide  ranges  of  habitat,  which  are 
indicated  elsewhere  (page  716).  They  are  readily  separable  from  other 
related  species  by  the  characters  outlined  on  pages  744  to  758.  The 
subfamilies  of  Tipulidae  may  be  separated  by  the  following  keys: 

Larvae 

1.  Body  provided  with  elongate  spines  or  leaflike  projections. 

Cylindrotominae,  pars  (p.  959) 
Body  without  distinct  spines 2 

2.  Form  depressed,  with  more  or  less  distinct  lateral  tubercles;  terrestrial  on  spermatophytic 

plants Cylindrotominae,  pars  (p.  959) 

Form  terete;  if  depressed,  without  tubercles 3 

3.  Spiracular  disk  surrounded  by  six  or  eight  lobes Tipulinae  (p.  974) 

Spiracular  disk  not  as  above 4 

4.  Spiracular  disk  surrounded  by  two,  four,  or  five  lobes 5 

Spiracular  disk  with  three  lobes  or  without  distinct  lobes 7 

5.  Head  capsule  massive,  the  hypopharynx  a  flattened  plate  with  few  teeth;  size  large 

(aberrant  Tipulinae) 6 

Head  capsule  massive  or  dissected,  if  the  former  the  hypopharynx  not  as  above;  size 
usually  small Limnobiinae,  pars  (p.  793) 

6.  Spiracular  disk  with  five  lobes;  lives  in  moss Genus  Dolichopeza  Curt.  (p.  981) 

Spiracular  disk  with  four  slender,  hornlike  lobes;  lives  in  earth. 

Tipula  selene  Meig.  (p.  1016) 

7.  Size  large  (30  mm.  or  over);  form  very  stout,  terete;  head  capsule  of  the  tipuline  type; 

lives  in  wood Genus  Tanyptera  Latr.  (p.  988) 

Size  small  (20  mm.  or  under);  form  slender,  terete;  head  capsule  of  the  limnobiine  type. 

Limnobiinae,  pars.  (p.  793) 

Pupae 

1.  Basal  abdominal  segments  unarmed  with  teeth  or  spinous  projections  before  posterior 

margin 2 

Basal  abdominal  segments  armed  with  a  transverse  row  of  usually  small  teeth  or  chitinous 
projections  before  posterior  margin 3 

2.  Last  larval  skin  adhering  to  posterior  end,  attaching  pupa  to  a  plant  stem  or  a  leaf; 

coloration  bright  green Cylindrotominae,  pars  (p.  959) 

Not  as  above Limnobiinae,  pars  (p.  793) 

3.  Maxillary  palpi  curved  or  recurved  at  tips;  size  large,  usually  12  mm.  or  over. 

Tipulinae,  pars  (p.  974) 
Maxillary  palpi  not  recurved  at  tips 4 

4.  Maxillary  palpi  long,  slightly  or  decidedly  curved  at  tips;  size  large,  length  usually  12  mm. 

or  over 5 

Maxillary  palpi  short,  straight;  size  small,  usually  10  mm.  or  under. 

Limnobiinae,  pars  (p.  793) 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  793 

5.  Dorsal  abdominal  segments  with  two  slender  spines  before  margin. 

Cylindrotominae,  pars  (p.  959) 
Dorsal  abdominal  segments  with  four  or  more  teeth  or  spines  before  margin. 

Tipulinae,  pars  (p.  974) 

SUBFAMILY  Limnobiimie 

The  subfamily  Limnobiinae  includes  a  vast  assemblage  of  usually 
small  crane-flies.  Only  a  few  genera  approach  the  ordinary  size  of  the 
other  principal  subfamily,  the  Tipulinae,  such  genera  being  Limnobia, 
Psaronius,  Limnophila,  Eriocera,  Pedicia,  and  a  few  others. 

The  writer  has  endeavored  to  key  the  immature  stages  of  tribes,  sub- 
tribes,  and  genera.  As  has  been  stated  elsewhere,  the  keys  are  based 
almost  entirely  on  material  seen  by  the  writer,  and  additional  specimens 
of  other  species  will  undoubtedly  modify  the  arrangement  very  con- 
siderably. It  is  believed,  however,  that  the  keys  as  given  will  at  least 
furnish  suggestions  or  a  basis  for  succeeding  work.  The  characters 
given  in  the  keys,  in  so  far  as  is  possible,  are  those  that  can  be  seen  with- 
out making  a  detailed  dissection  of  the  specimen.  However,  for  most 
species  it  is  necessary  to  study  the  larval  head,  as  already  outlined  (page 
741).  The  character  of  "  head  massive  "  or  "  head  rodlike  "  can  often 
be  detected  thru  the  larval  integument  without  dissection.  The  spi- 
racular  disk  is  usually  studied  without  especial  difficulty. 

The  immature  stages  of  the  majority  of  the  species  are  spent  in  moist 
earth,  usually  near  water.  Some  are  nearly,  if  not  quite,  aquatic  (Antocha, 
Elliptera,  some  Dicranomyia) ;  others  are  fungicolous  (Ula,  some  Lim- 
nobia); several  live  under  the  bark  of  trees  (some  Dicranomyia,  some 
Rhipidia,  Discobola,  Gnophomyia,  Teucholabis,  Elephantomyia,  and 
others);  one,  at  least,  mines  in  the  leaves  of  plants  (Dicranomyia). 

The  habits  of  the  various  genera  and,  species  are  discussed  in  greater 
detail  under  the  respective  titles.  The  following  keys  separate  the 
tribes  and  the  subtribes  of  the  subfamily  Limnobiinae: 

Larvae 

1.  Spiracular  disk  provided  with  two  long  ventral  lobes 2 

Spiracular  disk  not  as  above 3 

2.  Spiracles  lacking  or  vestigial;  mentum  not  completely  divided  medially;  hypopharynx  a 

chitinized  double  comb;  species  aquatic,  in  silken  cases. 

Limnobiini,  subtribe  Antocharia  (p.  799) 

Spiracles  large,  prominent,  exposed;  mentum  completely  divided  medially;  hypopharynx 
labriform Pediciini  (p.  894) 

3.  Spiracular  disk  surrounded  by  four  or  five  lobes r 4 

Spiracular  disk  surrounded  by  three  lobes  or  without  distinct  lobes 19 


794  CHAKLES  PAUL  ALEXANDER 

4.  Head  capsule  massive,  compact,  the  posterior  incisions  usually  shallow 5 

Head  capsule  of  four  or  six  slender  rods,  the  posterior  incisions  profound 15 

5.  Mentum  completely  divided,  a  toothed  plate  on  either  side;  abdominal  segments  without 

distinct  creeping-welts -6 

Mentum,  if  present  and  chitinized,  not  completely  divided;  abdominal  segments  with 
basal  creeping-welts 8 

6.  Spiracular  disk  squarely  truncated,  surrounded  by  five  lobes Eriopterini  (p.  908) 

Spiracular  disk  with  four  lobes 7 

7.  Each  mental  plate  four-toothed;  hypopharynx  labriform. 

Pediciini,  subtribe  Adelphomyaria  (p.  895) 
Each  mental  plate  with  seven  or  eight  teeth;  hypopharynx  a  comblike  ring. 

Hexatomini,  subtribe  Pseudolimnophilaria  (p.  848) 

8.  Spiracular  disk  with  five  lobes 9 

Spiracular  disk  with  four  lobes 12 

9.  Antennae  almost  globular,  with  two  conical  apical  papillae;  lives  in  fungi. 

Hexatomini,  subtribe  Ularia  (p.  838) 
Antennae  elongate-cylindrical 10 

10.  Abdomen  with  dorsal  and  ventral  creeping-welts;  mentum  with  more  than  five  teeth. 

Limnobiini  (p.  795) 
Abdomen  with  six  ventral  welts  only;  mentum  with  five  or  fewer  teeth 11 

11.  Mentum  five-toothed;  lives  in  earth Limnobiini,  subtribe  Rhamphidaria  (p.  830) 

Mentum  three-toothed;  lives  in  wood Hexatomini,  subtribe  Epiphragmaria  (p.  843) 

12.  Antennae  almost  globular,  with  two  conical  apical  papillae;  lives  in  fungi. 

Hexatomini,  subtribe  Ularia  (p.  838) 
Antennae  elongate-cylindrical 13 

13.  Abdomen  with  dorsal  and  ventral  creeping-welts,  the  latter  naked;  mentum  not  three- 

toothed;  forms  aquatic Limnobiini,  subtribe  Ellipteraria  (p.  806) 

Abdomen  with  ventral  creeping-welts  only;  mentum  with  only  three  primary  teeth; 
species  not  aquatic 14 

14.  Form  long,  slender;  skin  naked,  shiny,  transparent;  apical  segment  of  antennae  elongate, 

as  long  as,  or  longer  than,  basal  segment;  mentum  with  a  smaller  tooth  on  either  side; 

lives  in  earth Limnobiini,  subtribe  Dicranoptycharia  (p.  828) 

Form  short,  stout;  skin  white,  opaque;  apical  segment  of  antennae  short,  hemispherical; 
mentum  without  small  lateral  teeth;  lives  under  bark. 

Hexatomini,  subtribe  Epiphragmaria  (p.  843) 

15.  Blades  of  maxillae  not  produced;  form  long  and  slender 16 

Blades  of  maxillae  produced  into  flattened  elongate  appendages,  the  tips  of  which  pro- 
trude from  the  thoracic  orifice  when  the  head  is  completely  retracted;  form  short 
and  stout 17 

16.  Spiracular  disk  squarely  truncated,  surrounded  by  five  lobes  which  are  fringed  with 

numerous,  usually  short,   hairs;  esophageal  region  not  conspicuously  grooved. 

Eriopterini,  subtribe  Eriopteraria  (p.  911) 

Spiracular  disk  surrounded  by  four  lobes,  each  ventral  lobe  with  a  single  elongate  bristle; 
esophageal  region  elongate,  grooved;  lives  under  bark. 

Eriopterini,  subtribe  Elephantomyaria  (p.  952) 

17.  Mental  region  a  narrow,  transverse,  chitinized  bar. 

Hexatomini,  group  Limnophilae  (p.  858) 
Mental  region  not  chitinized 18 

18.  Mandibles  hinged;  maxillae  and  labrum  densely  hairy;  dorsal  plates  of  head,  capsule 

united  into  a  spatula Hexatomini,  group  Ulomorphae  (p.  869) 

Mandibles  not  hinged;  maxillae  and  labrum  not  densely  hairy;  dorsal  plates  of  head 
capsule  separated Hexatomini,  subtribe  Hexatomaria  (p.  876) 

19.  Head  capsule  massive,  compact 20 

Head  capsule  of  six  slender  rods;  lives  under  bark. 

Eriopterini,  subtribe  Eriopteraria  (p  911) 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  795 

20.  Mental  plates  not  completely  divided;  abdominal  segments  with  basal  creeping-welts 
on  both  ventral  and  dorsal  surface;   spiracular  disk  indistinctly  four-  or  five-lobed. 

Limnobiini  (p.  795) 

Mental  plates  completely  divided;  abdominal  segments  without  welts;  spiracular  disk 
obliquely  truncated Eriopterini  (p.  908) 

Pupae 

1.  Pronotal  breathing  horns  eight-branched;  forms  entirely  aquatic. 

Limnobiini,  sub  tribe  Antocharia  (p.  799) 
Pronotal  breathing  horns  simple,  unbranched 2 

2.  Rostral  sheath  elongated;  lives  in  wood.  .Eriopterini,  subtribe  Elephantomyaria  (p.  952) 
Rostral  sheath  not  elongated 3 

3.  Pronotal  breathing  horns  very  minute,  conical,  visible  only  with  a  lens.6 

Limnobiini,  subtribe  Dicranoptycharia  (p.  828) 
Pronotal  breathing  horns  larger,  not  microscopic 4 

4.  Dorsal  spiracles  on  eighth  abdominal  segment  large  and  functional 5 

Dorsal  spiracles  on  eighth  abdominal  segment  small  or  lacking 7 

5.  A  large  circular  spinous  area  on  abdominal  pleurites;  cephalic  crest  chitinized,  acutely 

pointed;  pronotal  breathing  horns  directed  ventrad;  lives  in  decaying  wood. 

Hexatomini,  subtribe  Epiphragmaria  (p.  843) 
Not  as  above 6 

6.  Pronotal   breathing  horns   large,  flattened,   the   tips  yellow;   abdominal   tergites  with 

shagreened  crossbands Hexatomini,  subtribe  Ularia  (p.  838) 

Pronotal  breathing  horns  slender,  cylindrical;  abdominal  tergites  with  tran verse  rows  of 
small  spines Limnobiini,  subtribe  Rhamphidaria  (p.  830) 

7.  Abdominal  pleurites  with  circular  areas  set  with  numerous  microscopic  spicules;  pronotal 

breathing  horns  short,  usually  truncated  at  tips,  which  are  margined  with  the  breathing 

pores Pediciini  (p.  894) 

Abdominal  pleurites  not  as  above,  if  with  spines  these  large  and  few  in  number;  pronotal 
breathing  horns  long,  cylindrical 8 

8.  Abdominal  segments  with  broad  transverse  bands  or  welts  on  basal  rings  of  third  to 

seventh  tergites Limnobiini  (p.  795) 

Abdominal  segments  with  basal  ring  unarmed  as  above,  posterior  ring  before  margin 
with  a  transverse  row  of  spines  or  stiff  setae 9 

9.  A  distinct  crest  on  mesonotal  prescutum  armed  with  tubercles,  spines,  or  setae;  size 

small  (usually  under  9  mm.) Eriopterini  (p.  908) 

No  distinct  crest  on  mesonotal  prescutum  (scutellum  armed  in  some  Eriocera);  size 
large  (usually  over  10  mm.) 10 

10.  Leg  sheaths  very  short,  barely  exceeding  wings;  lives  under  bark. 

Eriopterini,  genus  Gnophomyia  (p.  934) 
Leg  sheaths  longer,  extending  one  or  more  segments  beyond  tips  of  wings 11 

11.  Size  small  (under  6  mm.);  abdominal  armature  weak,  lacking  on  segment  7. 

Pediciini,  subtribe  Adelphomyaria  (p.  895) 

Size  larger;  abdominal  armature  stronger,  spinous;  if  small  in  size  (Dicranophragma), 
basal  annuli  of  abdominal  segments  armed  with  naked  tubercles . .  Hexatomini  (p.  835) 

Tribe  Limnobiini 

A  large  group  of   crane-flies,  arranged  in  a  few  often  extensive  genera, 
comprise  the  tribe  Limnobiini.     At  first  sight  the  tribe  appears  to  be  a 

5 In  the  genus  Cladura  (Eriopterini),  reared  while  this  paper  was  going  thru  the  press,  the  breathing 
pores  are  likewise  microscopic,  being  entirely  sessile  (page  949). 


796  CHARLES  PAUL  ALEXANDER 

heterogeneous  assemblage,  but  in  reality  it  constitutes  a  natural  group. 
The  tribe  as  herein  arranged  includes  the  old  group  Limnobiini,  with  the 
addition  of  several  genera  that  were  formerly  distributed  in  the  Antochini. 
The -divisions  of  the  tribe  as  now  constituted  are  as  follows: 

1.  Limnobaria  —  including  the  old  tribe  Limnobiini. 

2.  Ellipteraria  —  including  the  genus  Elliptera.     This  is  close  to  the 
preceding  subtribe  and  may  be  a  group  belonging  to  it. 

3.  Antocharia  —  including  Antocha  and  presumably  allied  genera,  as 
Orimargula,  Orimarga,  Diotrepha,  and  possibly  others. 

4.  Rhamphidaria  —  including  Rhamphidia  and  its  allies. 

5.  Dicranoptycharia  —  including   Dicranoptycha   only. 

These  groups  are  not  far  removed,  phylogenetically,  from  the  lowermost 
divisions  of  the  Hexatomini,  such  as  the  Ularia  and  the  Epiphragmaria, 
and  the  two  tribes  are  unquestionably  closer  together  than  their  arrange- 
ment on  paper  would  indicate.  The  separation  of  the  two  major  groups 
was  made  largely  on  the  characters  of  the  imagines. 

The  larvae  have  the  body  terete,  moderately  elongate  or  very  long  and 
slender  (Dicranoptycha).  The  abdominal  segments  are  subdivided  into 
a  basal  and  a  posterior  ring,  the  former  with  transverse  welts  of  micro- 
scopic chitinized  points  or  hooks.  In  the  Limnobaria  and  the  Antocharia 
these  welts  occur  on  both  the  dorsal  and  ventral  surfaces  in  the  form  of 
microscopic  hooks;  in  the  Ellipteraria  they  are  on  segments  3  to  9  on  the 
dorsal  surface  only,  being  indicated  on  the  ventral  surface  but  naked; 
in  the  Rhamphidaria  the  welts  are  ventral  in  position  on  segments  2  to 
7;  in  the  Dicranoptycharia  they  are  similar,  on  segments  2  to  8.  The 
body  in  Dicranoptycha  is  entirely  glabrous. 

The  head  capsule  is  of  moderate  to  large  size  and  is  massive  and  com- 
pact, consisting  of  a  narrow  dorsal  plate  which  is  usually  indented  behind, 
arid  two  broad  mussel-shaped  lateral  plates  which  are  connected  ante- 
riorly across  the  venter  to  form  the  mental  plate.  The  mental  plate 
consists  of  an  outer  plate  which  usually  terminates  in  a  single  median 
point,  and  behind  this  an  inner  plate  which  contributes  additional  teeth 
to  the  mentum.  In  Dicranoptycha  there  is  but  one  subequal  tooth  on 
either  side,  with  an  additional  much-reduced  tooth;  in  Rhamphidia 
there  are  two  teeth,  and  in  the  other  groups  there  are  usually  four  or  five. 
The  hypopharynx  is  usually  a  double  plate  united  at  the  ends  to  form 
a  collar,  with  the  anterior  margins  finely  toothed.  The  maxillae  are 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  797 

large  and  simple  in  structure;  the  cardo  and  stipes  are  large;  the  palpus  is 
flattened.  The  antennae  have  the  apical  segment  or  papilla  ranging  from 
elongate,  in  Dicranoptycha,  to  very  flattened  and  disklike,  in  Limnobia 
and  its  allies.  The  mandibles  are  usually  of  simple  structure,  with  one  or 
two  dorsal  teeth  and  from  three  to  seven  teeth  in  the  ventral  cutting 
row. 

The  spiracles  are  lacking  in  some  species,  at  least,  of  Antocha.  The 
spiracular  disk  is  surrounded  in  Rhamphidia  by  five  subequal  lobes,  in 
Dicranoptycha  by  four  slender,  naked  lobes;  in  many  Limnobaria  the  lobes 
are  lacking  or  indistinct. 

The  larvae  of  many  of  the  species  are  able  to  spin  silken  cocoons  or 
tubes  in  which  they  live.  These  tubes  are  open  at  both  ends,  and  are 
usually  covered  exteriorly  with  particles  of  extraneous  matter  gathered 
in  the  larval  haunts. 

The  pupae  usually  lack  a  distinct  setiferous  cephalic  crest,  altho  one 
is  present  in  Rhamphidia  and  in  Dicranoptycha.  The  pronotal  breathing 
horns  are  usually  large,  and  are  either  subcircular,  or  wider  than  long  (most 
Limnobaria),  rarely  elongate  (Rhamphidaria),  very  large,  earlike,  and 
contiguous  or  practically  so  on  the  median  line  (Ellipteraria),  or  branched 
into  eight  long  filaments  (Antocharia) ;  in  the  Dicranoptycharia,  how- 
ever, they  are  microscopic.  The  abdominal  segments  on  the  basal  ring 
often  show  a  transverse  welt  of  small  hairs  or  a  double  convergent  row  of 
chitinized  hooks;  in  Discobola,  Rhamphidia,  and  Dicranoptycha,  how- 
ever, this  is  apparently  not  the  case,  the  abdominal  armature  being  more 
eriopterine  or  hexatomine  in  appearance.  The  eighth  abdominal  seg- 
ment often  bears  a  pair  of  dorsal  spiracles;  these  are  apparently  lacking 
in  some  species  (Antocha  saxicola)  and  are  small  in  most  Limnobaria,  but 
are  large  and  functional  in  Rhamphidia. 

The  following  keys  separate  the  subtribes  of  the  tribe  Limnobiini: 

Larvae 

1.  Body  ending  in  two  long  vertral  lobes;  spiracles  lacking  or  very  reduced;  forms  strictly 

aquatic Antocharia  (p.  799) 

Body  not  as  above;  spiracles  large 2 

2.  Body  with  ventral  and  dorsal  welts  on  abdominal  segments 3 

Body  with  ventral  welts  only 4 

3.  Spiracular  disk  surrounded  by  four  lobes  which  are  provided  with  long  fringes  of  hair; 

dorsal  welts  microscopically  spiculose;  ventral  welts  naked;  species  aquatic. 

Ellipteraria  (p.  806) 
Spiracular  disk  not  as  above;  dorsal  and  ventral  welts  alike Limnobaria  (p.  808) 


798  CHARLES  PAUL  ALEXANDER 

4.  Body  moderately  elongated,  covered  with  a  long,  dark  pubescence;  abdomen  squarely 
truncated  at  end,  surrounded  by  five  lobes,   presenting  an  eriopterine  appearance; 

mentum  conspicuously  five-toothed Rhamphidaria  (p.  830) 

Body  very  long  and  slender,  glabrous;  abdomen  surrounded  by  four  narrow,  glabrous 
lobes;  mentum  indistinctly  five-toothed Dicranoptycharia  (p.  828) 

Pupae 

1.  Pronotal  breathing  horns  branched;  forms  entirely  aquatic Antocharia  (p.  799) 

Pronotal  breathing  horns  simple 2 

2.  Pronotal  breathing  horns  very  tiny,  microscopic,  conical Dicranoptycharia  (p.  828) 

Pronotal  breathing  horns  large,  conspicuous 3 

3.  Pronotal  breathing  horns  large,  earlike,   contiguous  basally;  forms  aquatic,  in  silken 

cocoons Ellipteraria  (p.  806) 

Pronotal  breathing  horns  not  contiguous  basally 4 

4.  Cephalic  crest  small  or  lacking;  pronotal  breathing  horns  short  and  broad,  rarely  elongated; 

a  pair  of  small  spiracles  on  dorsum  of  eighth  abdominal  segment. .  .Limnobaria  (p.  808) 
Cephalic  crest  large,  setiferous;  pronotal  breathing  horns  long  and  slender,  cylindrical; 
a  pair  of  large  spiracles  on  dorsum  of  eighth  abdominal  segment. 

Rhamphidaria  (p.  830) 

The  most  important  literature  on  the  tribe  Limnobiini  is  as  follows : 

Antocha  saxicola General Needham,  1908a:  205. 

Antocha  sp Larva Malloch,  1915-17  b:  236-237. 

Elliptera  omissa Larva,  pupa,  general. .  .  Mik,  1886  b. 

Elliptera  omissa Larva,  pupa Griinberg,  1910:31-32.    (Copy.) 

Elliptera  omissa Larva,  pupa Malloch,    19 15-17 b: 226-227. 

Thaumastoptera  calcezti Larva,  pupa,  general.  .  .   Lenz,  1920a. 

Limnobia  quadrimaculata General Von  Roser,  1834  (as  annulus). 

Limnobia  quadrimaculata Larva,  pupa Beling,  1873b:  590-591  (as  annu- 
lus}. 

Limnobia  bifasciata Larva,  pupa,  general. .  ..  Bremi-Wolf,    1846  (as   xanthop- 

terd). 

Limnobia  bifasciata Larva,  pupa,  general.  .  .   Paste jrfk,  1909  (as  xanthopterd) . 

Limnobia  bifasciata Larva,  pupa De  Meijere,  1916: 198-201. 

Limnobia  decemmaculata General Loew>  1873:41. 

Limnobia  decemmaculata General Verrall,  1912. 

Limnobia  flavipes Larva. Beling,  1886: 202. 

Limnobia  inuskt General Beling,  1886: 202    (as  macro- 
stigma). 

Limnobia  sexpunctata Larva Beling,  1879:54-55  (as  nigro- 

punctata). 

Limnobia  nubeculosa General Beling,  1879:56. 

Limnobia  obscuricornis. . Larva,  pupa Beling,  1879: 55-56. 

Limnobia  tripunctata Larva,  pupa Beling,  1873 b: 591-592. 

Limnobia  triocellata Larva,  pupa,  general.  .  .   Johnson,  1906:2. 

Limnobia  triocellata Larva,  pupa Malloch,  19 15-17 b: 215-216. 

Limnobia  immatura Pupa. . Malloch,  1915-17 b:  216. 

Libnotes  perkinsi General Perkins,  1913:  clxxxii  (as  Limno- 
bia). 

Discobola  caesarea Pupa Mik,  1884. 

Dicranomyia  trinotata Larva,  pupa Thienemann,  1909. 

Dicranomyia  trinotata Larva,  pupa Griinberg,  1910:29.     (Copy.) 

Dicranomyia  dumetorum General Winnertz,  1853. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  799 

Dicranomyia  dumetorum General Beling,  1873  b:  592. 

Dicranomyia  dumetorum General Beling,  1879:56. 


Dicranomyia  dumetorum Larva. 

Dicranomyia  sp General. 

Dicranomyia  foliocuniculator General. 

Dicranomyia  foliocuniculator Larva,  pupa. 


Beling,  1886:201-202. 
Schubart,  1854. 
Swezey,  1913. 
Swezey,  1915:87. 


Dicranomyia  umbrata Larva De  Meijere,  1916: 197-198. 

Dicranomyia  simulans Larva,  pupa,  general.  .  .  Needham,  1908 a: 214-217. 

Dicranomyia  simulans Larva,  pupa Malloch,  19 15-17 b: 213-214. 

Rhipidia  maculata Pupa Beling,  1873  b :  592. 

Rhipidia  maculata Larva,  general Beling,  1879:52-53. 

Rhipidia  uniseriata Larva,  general Beling,  1879:53-54. 

Rhipidia  domestica General Johnson,  1910:704. 

Dicranoptycha  winnemana Larva,  pupa Alexander,  1919  b. 

Rhamphidia  longirostris General Gercke,  1884. 

Rhamphidia  longirostris General Griinberg,  1910:30.     (Copy.) 

Rhamphidia  flavipes Larva Hart,  1898  [1895] :  197-199. 

Rhamphidia  flavipes Larva Malloch,  19 15-17 b: 231-232. 

Subtribe  Antocharia 

The  subtribe  Antocharia  includes  the  genus  Antocha  and  probably 
three  or  four  related  genera,  such  as  Diotrepha,  Orimarga,  and  Orimargula. 
The  group  is  well-defined  in  all  stages,  so  far  as  these  are  known,  the 
larvae  presenting  a  curious  superficial  resemblance  to  those  of  Pedicaria, 
while  the  pupae  introduce  a  novelty  of  structure  of  the  breathing  horns, 
which  is  discussed  in  detail  elsewhere,  (page  805).  It  is  probable,  however, 
that  these  peculiarities  of  larval  and  pupal  structure  are  largely  the  result 
of  habit  and  habitat,  and  a  critical  survey  of  the  structure  shows  a  close 
relationship  with  the  other  subtribes  herein  recognized. 

Genus  Antocha  Osten  Sacken  (Gr.  close  approximation) 
1859    Antocha  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  219. 

Larva. —  Body  slender,  tapering  behind,  ending  caudally  in  two  elongate  ventral  lobes 
which  bear  a  few  hairs  at  their  tips  and  at  intervals  along  their  length.  Abdominal  segments 
2  to  7  each  with  a  swollen  area  on  basal  ring  densely  covered  with  microscopic  hairs. 
Tracheal  gills  four  in  number,  large,  constricted  into  three  or  four  lobes.  Spiracles  lacking 
or  rudimentary.  Head  capsule  moderate  in  size.  Mentum  with  nine  or  ten  teeth,  deeply 
split  behind.  Maxilla  conspicuous,  consisting  of  two  subequal  lobes  which  are  provided  with 
dense  brushes  of  hairs.  Hypopharynx  with  chitinized  teeth. 

Pupa. —  Anterior  end  of  body  large,  tapering  behind.  Head  with  a  small  median  lobe 
in  front,  on  either  side  of  which  is  a  small  tubercle;  genae  gibbous.  Pronotal  breathing 
horns  large,  flattened,  the  margin  branching  into  eight  long  filaments.  Abdominal  segments 
on  basal  ring  with  a  double  transverse  row  of  small  hooks  which  converge  at  the  ends  to  inclose 
an  oval  depressed  area;  last  segment  of  body  terminating  in  two  strong,  recurved,  chitinized 
hooks. 


800  CHARLES  PAUL  ALEXANDER 

Antocha  is  a  small  genus  of  crane-flies  (about  seven  species)  whose 
specific  limits  are  still  not  well  understood.  The  species  are  well  dis- 
tributed thruout  the  Northern  Hemisphere.  The  adult  flies  are  of  prim- 
itive organization,  but  the  larvae  and  the  pupae  are  highly  specialized 
in  many  respects. 

The  only  previous  record  of  the  immature  stages  of  any  member  of  this 
group  is  the  unknown  Limnobiine  No.  2  (Malloch,  191 5-17 b :  236-237) ,  which 
surely  refers  to  an  Antocha,  possibly  A.  monticola  Alex.  The  main  point 
of  difference  between  the  species  described  by  Malloch  and  the  species 
described  in  detail  hereinafter  is  the  small  spiracles  mentioned  in  the 
description  of  the  former  species.  A.  saxicola  lacks  spiracles,  since  it 
has  no  use  for  them,  being  confined  to  submerged  cases  often  many  feet 
below  the  surface  of  the  water. 

Altho  nothing  is  known  concerning  the  immature  stages  of  the  genera 
Orimargula,  Orimarga,  and  Diotrepha,  the  writer  believes  that  these 
genera  will  be  found  to  have  larvae  of  this  same  general  type,  since  from 
the  structure  of  the  adults  they  are  obviously  derived  from  the  Antocha 
stem.  The  larvae  are  curiously  suggestive  of  the  Pediciini  (as  compared 
with  Dicranota),  but  the  structure  of  the  mouth  parts,  the  smooth  pseudo- 
pods,  and  the  cauda,  are  quite  different  and  indicate  that  the  similarities 
are  analogous  only.  The  pupae  are  unlike  those  of  any  crane-fly  as  yet 
made  known,  in  the  very  remarkable  breathing  horns;  but  this  is  possibly 
a  condition  brought  about  by  the  habitat  rather  than  a  fundamental 
feature.  The  apparently  very  different  pupae  of  Elliptera  are  closely 
related  to  Antocha. 

Antocha  saxicola  O.  S. 

1859  Antocha  saxicola  0.  S.  Proc.  Acad.  Nat.  Sci.  Phila.,  p.  219. 
Antocha  saxicola  has  one  of  the  most  interesting  life  histories  of  any  of 
the  crane-flies  yet  discovered,  not  only  because  of  the  larval  and  pupal 
habitat,  but  also  because  of  the  peculiar  structures  that  appear  in  the 
larva  and  in  the  pupa  and  have  been  found  nowhere  else  in  the  immature 
stages  of  the  family,  so  far  as  is  known.  The  larvae  simulate  strikingly 
the  same  stage  in  the  Pedicaria,  but  are  apneustic,  entirely  lacking  func- 
tional spiracles  and  depending  wholly  on  tracheal  gills  for  their  respira- 
tion. The  pupae  have  the  pronotal  breathing  horns  split  into  eight  long 
filaments,  so  that  they  bear  a  curious  superficial  reseml  lance  to  the  pupae 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  801 

of  the  black  fly  (Simuliidae).  Both  larvae  and  pupae  spend  their  entire 
lives  in  cases  on  stones  in  water  —  usually  in  running,  well-aerated  water, 
and  often  in  the  most  rushing  torrents. 

The  larvae,  &s  already  stated,  lack  spiracles,  the  entire  respiration  being 
carried  on  thru  tracheal  gills,  four  in  number,  and  the  rich  tracheal 
development  in  the  elongate  caudal  lobes.  Thus  the  tracheal  system  is 
truly  closed,  and  represents  the  maximum  of  specialization  in  the  reduc- 
tion in  size  and  final  loss  of  the  spiracles.  In  air-breathing,  terrestrial 
forms,  the  spiracles  are  large  and  situated  comparatively  close  together, 
gradually  becoming  smaller  and  more  removed  from  one  another  as  the 
creature  becomes  more  and  more  dependent  on  blood  gills  or  tracheal 
gills  for  respiration.  This  is  the  only  truly  closed  tracheal  system  known 
to  the  writer  to  occur  in  the  Tipulidae.  It  should  be  noted  that  the  loss 
of  the  spiracles  is  accompanied  by  great  enlargement  of  the  gills  and  the 
taking  on  of  the  gill  function  by  the  two  caudal  lobes.  The  haunt  of  the 
larvae  is  in  silken  cases  on  rocks,  often  in  the  swiftest  part  of  the  stream, 
where  it  is  impossible  to  breathe  thru  spiracles  and  where  the  associated 
forms  of  life  (Ephemerida,  Plecoptera,  Trichoptera,  and  Diptera)  all, 
or  practically  all,  depend  entirely  on  gills  for  .respiration. 

As  a  rule,  the  larval  cases  are  made  on  rubble  or  rounded  stones,  a 
crevice  or  a  groove  caused  by  inequalities  of  the  rock  surface  covered 
over  being  the  simplest  and  commonest  place  chosen.  The  inequalities 
in  the  rock  are  bridged  over  by  a  silken,  mud-  or  silt-covered  case,  which 
is  very  delicate  and  laterally  fimbriated  with  the  young  larva  but  becomes 
much  firmer,  thicker,  and  more  compact  with  the  older  larva  and 
pupa.  The  larval  case  is  open  at  both  ends  and  the  larva  passes  back- 
ward and  forward  freely,  showing  considerable  agility  when  disturbed. 
When  the  larva  is  still  small,  the  case  is  correspondingly  small  and 
insignificant;  but  the  case  of  the  matured  larva  is  conspicuous,  measur- 
ing from  4  to  5  centimeters  in  length  and  about  1.2  centimeters  across 
the  lateral  "  wings,"  or  fimbriations.  The  insect  moves  freely  along 
the  tube  but  is  very  loath  to  leave  it  unless  actually  ejected.  As 
stated  above,  the  late  larval  and  pupal  covering  is  very  different  from  the 
flimsy,  silt-covered  tube  of  the  young  larva,  being  smooth,  compact, 
hard,  and  often  covered  with  pebbles.  The  pupa  has  two  powerful  hooks 
at  its  caudal  end,  enabling  it  to  fasten  to  the  case.  In  most  cases  the  pupa 
hangs  with  the  current,  head  downstream,  like  the  pupa  of  Blepharocera 


802  CHARLES  PAUL  ALEXANDER 

and  unlike  the  somewhat  similar-appearing  pupa  of  the  Simuliidae,  which 
rests  with  the  head  upstream,  against  the  current,  the  pupal  case  being 
open  at  the  cephalic  end  only. 

These  curious  larvae  were  first  noted  at  Ithaca,  New  York,  in  mid- 
April,  in  Cascadilla  Creek.  They  were  common  in  situations  such  as 
described  above.  Larvae  were  found  thruout  most  of  April,  May,  and 
June;  they  would  probably  be  seen  in  somewhat  fewer  numbers  thru 
most  of  the  summer  season,  since  the  adult  flies  have  a  long  seasonal 
appearance.  The  first  pupa  was  found  on  May  15,  1917,  but  the  season 
that  year  was  very  backward  and  undoubtedly  the  species  pupates  earlier 
in  more  nearly  normal  seasons. 

In  Cascadilla  and  Fall  Creeks,  at  Ithaca,  the  immature  stages  of  Antocha 
are  usually  associated  with  a  fauna  of  rapid-water  (lotic)  forms,  the 
following  being  the  more  notable  and  constant: 

Planarians.          Planaria  sp. 

Ephemeridae.  Nymphs  of  Baetis,  Leptophlebia,  Ephemerella,  Ecdyurus, 
-Epeorus,  Iron,  Heptagenia,  Chirotenetes,  and  others. 

Perlidae.  Nymphs  of  Pteronarcys,  Perla  immarginata  Say,  Acro- 

neuria,  Neoperla,  and  others. 

Trichoptera.  Larvae  and  pupae  of  Helicopsyche  (abundant),  Hydro- 
psyche,  Hydropsychodes,  Ithytrichia,  Rhyacophila,  Lep- 
tocerus,  Poly  cent  ropus,  and  others. 

Lepidoptera.       Larvae  and  pupae  of  Elophila. 

Coleoptera.         Larvae  of  Psephenus. 

Diptera.  Larvae  and  pupae  of  Blepharocera,  Simulium,  Ortho- 

cladius,  Tanytarsus,  and  others. 

Early  hi  spring  the  rocks  are  plastered  with  dense  coatings  of  Diato- 
maceae  (Navicula,  Synedra,  Meridion,  and  other  genera),  which  later 
in  the  season  become  much  rarer  or  disappear  entirely.  It  is  often  impos- 
sible to  tell  the  case  of  Antocha  from  that  of  some  very  similar  caddis- 
worm  cases,  especially  some  of  the  glossosomatine  Rhyacophilidae.  Dr. 
Noyes  found  larvae  of  Antocha  in  a  small,  rapid-flowing  stream  neai 
Ringwood  Hollow,  and  here  the  cases  were  covered  with  tiny  pebbles 
and  it  was  quite  impossible  to  distinguish  them  superficially  from  asso- 
ciated caddis- worm  cases.  In  Cascadilla  Creek  the  little  cases  of  Heli- 
copsyche often  plaster  the  upper  surfaces  of  submerged  rocks,  and  the 
writer  has  found  tubes,  of  Antocha  that  were  almost  buried  beneath  these 
cases. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  803 

Antocha  is  by  no  means  confined  to  rapidly  flowing  streams,  altho 
the  insects  are  very  often  found  in  such  situations.  The  writer  has  found 
adults  in  his  tent  traps  set  over  the  Sacandaga  River,  in  Fulton  County, 
New  York,  where  the  water  was  very  quiet  and  at  least  ten  feet  deep. 
Needham  (1908 a:  169-170,  205),  similarly,  found  adults  in  numbers  in 
his  tent  traps  set  over  Beaver  Meadow  Brook  at  Old  Forge,  New  York, 
in  August,  1905,  but  did  not  locate  the  larvae.  In  rapid-flowing  streams 
the  insects  sometimes  occur  just  at  the  surface  in  a  few  millimeters  of 
water,  or  at  greater  depths.  The  immature  stages  seem  adapted  to  live 
under  almost  any  conditions  of  current,  from  moderate  pressure  to  sit- 
uations where  the  water  rushes  by  in  torrents  and  where  but  few  of  the 
usual  lotic  organisms,  such  as  Blepharocera,  Simulium,  Psephenus,  and 
others,  can  exist.  From  Clemens'  studies  (1917:14-23)  it  is  evident  that 
the  current  is  much  more  rapid  just  beneath  the  surface  than  at  various 
lower  depths,  so  that  at  a  depth  of  one  foot  the  current  velocity  is  only 
about  two-thirds  of  that  at  the  surface.  Thus  these  aquatic  organisms  are 
not  constantly  and  entirely  subjected  to  such  tremendous  pressures  as 
on  first  sight  they  appear  to  be.  Many  of  the  larvae  and  pupae  perish 
from  desiccation,  due  to  the  lowering  of  the  stream  level  and  the  conse- 
quent exposure  of  the  rocks  on  which  their  homes  are  made. 

Copulation  between  the  adult  flies  takes  place  on  the  exposed  rocks 
in  and  along  the  margins  of  the  streams  where  the  larvae  live  (Osten 
Sacken,  1869:127).  The  eggs  are  deposited  in  the  water,  and  the  entire 
life,  until  the  emergence  of  the  adult  fly,  is  spent  beneath  the  water.  The 
whole  life  cycle  may  require  a  year,  altho  the  species  is  possibly  double- 
brooded.  At  a  single  time,  and  even  on  a  single  rock,  larvae  of  various 
sizes,  from  very  small  ones  to  those  almost  fully  grown,  may  be  found, 
and  this  probably  explains  the  long  flight-period  of  the  adult.  That  the 
species  is  double-brooded  remains  to  be  proved. 

The  food  of  the  larva  consists  of  microscopic  plant  organisms  in  the 
water,  the  curious  maxillae,  with  their  dense  brushes  of  long  hairs, 
undoubtedly  being  an  adaptation  for  this  type  of  food. 

Larva.—  Total  length,  9.5-10.5  mm. 

Length  of  caudal  lobes  alone,  1.1-1.2  mm. 
Diameter  of  body,  1-1.1  mm. 

Coloration  of  living  larva,  light  greenish  brown  above,  clearer  greenish  ventra]ly;  contents 
of  alimentary  canal  showing  clearly  thru  the  thin  skin;  on  segments  7  and  8,  two  paired,  bright 


804  CHARLES  PAUL  ALEXANDER 

orange  bodies  which  are  very  conspicuous,  these  possibly  being  fatty  in  nature;  welts  on  the 
abdomen  dark  brown;  in  preserved  specimens  general  coloration  fading  to  a  dull  yellow. 

Form  elongate  (Plate  XX,  57),  tapering  behind.  Surface  with  a  dense,  appressed  pubes- 
cence and  scattered  erect  hairs.  Prothorax  long,  narrowed  in  front,  anterior  orifice  margined 
with  dense,  fine  pubescence;  sides  of  prothorax  with  numerous  long,  erect,  pale  hairs. 
Mesothorax  and  metathorax  indistinctly  divided  into  two  approximately  equal  annuli; 
anterior  annulus  with  a  few  lateral  setae.  First  abdominal  segment  short,  the  setae  arranged 
as  follows:  dorsal,  two  small  grouped  setae  near  posterior  margin,  laterad  of  each  of  these, 
but  in  alinement,  a  solitary  stout  seta,  proximad  of  each  of  these  a  stout  seta,  and  cephalad 
of  each  of  these  another  strong  seta,  the  principal  setae  of  each  side  thus  forming  a  rough 
triangle;  a  long,  delicate  pleural  seta  on  each  side;  ventral,  two  small  groups  of  setae,  with 
an  arrangement  similar  to  that  of  the  grouped  setae  of  dorsum.  Abdominal  segments  2  to  7 
each  indistinctly  divided  into  two  annuli  by  a  transverse  constriction,  the  anterior  ring  about 
half  the'length  of  the  posterior  ring  and  bearing  medially  a  transverse  elongate-oval  (dorsal) 
to  short-oval  (ventral)  welt,  covered  with  microscopic  points;  ventral  welts  very  convex 
and  swollen;  pleura  with  a  long  seta;  posterior  ring  with  setae  arranged  as  described  above 
for  first  abdominal  segment;  setae  of  posterior  segments  of  body  longer,  but  occupying  same 
relative  position;  small  solitary  inner  seta  of  dorsal  posterior  line  becoming  large  and  prom- 
inent on  seventh  segment; -segment  8  with  six  powerful  setae  in  alinement  on  ventral  surface, 
situated  at  base  of  gills,  two  being  pleural  and  four  ventral  in  position;  dorsum  of  segment 
8  with  a  rounded  median  lobe.  Gills  four,  long,  delicate,  divided  into  lobes  by  constrictions 
(the  two  caudal  lobes,  as  well  as  the  gills,  have  taken  on  a  respiratory  function).  Caudal 
ventral  lobes  two  (Plate  XXI,  68)  very  long,  parallel,  with  scattered  setae  arranged  as  fol- 
lows: at  tips,  six  or  seven;  at  about  two-thirds  length,  three;  at  about  one-third  length,  a  tuft 
of  from  fifteen  to  twenty  long  and  short  setae  on  lateral  and  dorsal  faces;  a  small  solitary  seta, 
dorsal  in  position,  at  base  of  lobe;  a  similar  bristle  on  ventral  face  at  about  midlength  of  lobe. 
Like  remainder  of  body,  cauda  covered  with  an  abundance  of  delicate  appressed  pubescence; 
on  dorsal  side,  at  base  of  each  lobe,  a  rounded  spot,  and  just  inside  this  a  narrow,  longitudinal 
line  which  is  destitute  of  pubescence.  Only  dorsal  lobe  the  median  one  of  eighth  segment, 
mentioned  above.  Spiracles  lacking. 

Head  capsule  (Plate  XX,  58)  moderate  in  size.  Lateral  plates  thin,  double,  inner  one  the 
longest,  outer  one  forming  mental  plate.  Dorsal  plate  broad  in  front,  narrowed  behind, 
posterior  margin  bluntly  notched;  in  front  of  this  plate,  two  broad  plates,  rather  widely 
separated  medially,  presumably  belonging  to  clypeus.  Labrum  broad,  cephalic  margin  and 
ventral  face  with  transverse  rows  of  short  hairs.  Mentum  (Plate  XX,  59)  conspicuous,  deeply 
split  behind  but  not  entirely  divided  as  in  the  Pediciini;  an  outer  flattened,  circular  median 
lobe  whose  outer  face  is  covered  with  small,  scalelike  roughenings;  behind  (dorsad  of)  this 
outer  lobe  the  mentum  proper,  roughly '  triangular  in  outline,  conspicuous,  margin  with  a 
broad,  blunt,  median  tooth  which  is  sometimes  (Plate  XXI,  66)  bilobed  to  form  two  subequal 
apical  teeth;  besides  this  median  tooth,  four  lateral  teeth,  the  outermost  one  broad  with  its 
lateral  angle  rounded.  Hypopharynx  (Plate  XXI,  64)  forming  a  ring  into  which  ducts  of  sali- 
vary glands  open;  anterior  part,  somewhat  resembling  mentum  in  shape,  a  narrow  blade 
with  anterior  margin  having  about  eight  teeth;  posterior  part  a  transverse,  arcuated  band 
with  anterior  margin  having  about  twenty  teeth.  (In  the  figure,  the  two  parts  of  the  hypo- 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  805 

pharynx  are  shown  diagrammatically  and  separated;  in  some  specimens  the  lateral  teeth  are 
more  acute,  in  others  they  are  more  rounded.)  Antenna  (Plate  XXI,  65)  elongate,  cylindrical, 
chitinized,  apex  pale,  with  two  long,  sensory  setae  and  a  few  papillae.  Mandible  (Plate  XXI, 
63  and  67)  strong,  flattened,  with  two  powerful  bristles  on  back,  or  scrobal  region,  near 
base;  inner  face  concave,  tip  ending  in  a  long  tooth,  dorsad  of  apex  a  single  smaller  tooth, 
ventral  cutting  edge  with  four  gradually  smaller  teeth,  beyond  the  last  of  which  the  margin 
is  crenulated  into  four  or  five  indistinct  carunculations;  viewed  from  inside,  lateral  teeth 
appearing  blunt.  Maxilla  (Plate  XXI,  63)  large,  consisting  of  two  elongate-oval  lobes,  the 
inner  one  densely  hairy;  palpus,  borne  at  tip  of  outer  lobe  on  ventral  face,  shaped  like  one- 
half  of  a  cylinder  split  lengthwise,  several  tiny  hyaline  sense  pegs  at  apex;  laterad  of  palpus 
and  nearer  base  of  outer  lobe,  a  small  elongate  sensory  tubercle  with  hairs  at  apex;  inner 
lobe  of  maxilla  subequal  in  size  and  length  to  outer  lobe,  but  more  densely  hairy;  on  its  ventral 
face,  four  or  five  long  sensory  tubercles  which  are  expanded  at  their  ends  into  setiferous  heads; 
at  base  of  maxilla,  a  long,  slender  arm  with  three  setiferous  punctures  at  apex  and  another 
puncture  at  about  two-thirds  length;  setae  of  this  arm  very  long  and  delicate.  (A  dorsal 
view  of  the  larva  is  shown  in  Plate  XX,  57.) 

Pupa. —  Length  to  tip  of  cephalic  crest,  6.2-6.8  mm. 
Width,  d.-s.,  l'.4-1.5  mm. 
Depth,  d.-v.,  1.1-1.2  mm. 

Head,  thorax,  and  sheaths  of  appendages  dark  brown  in  fully  colored  individuals;  abdomen 
pale  yellowish  white;  terminal  hooks  of  abdomen  heavily  chitinized. 

Head  on  margin  above  eyes  with  a  blunt  median  lobe  and  on  either  side  a  small  but  prom- 
inent tubercle;  gena  gibbous.  Compound  eyes  large,  semicircular  in  outline.  Front  between 
eyes  with  margins  almost  parallel.  Labrum  with  apex  truncated  or  indistinctly  bilobed. 
Labial  lobes  broad,  appearing  subtriangular.  Sheaths  of  maxillary  palpi  not  greatly 
elongated,  slender,  cylindrical. 

Pronotal  breathing  horns  (Plate  XX,  61)  flattened  at  base,  each  arcuated  basally  behind, 
bending  laterad  to  form  a  concave  hollow  in  front;  base  dark  brown,  chitinized,  branched 
into  eight  long,  pale  filaments  which  are  grouped  more  or  less  in  pairs;  the  two  ventral  and 
the  four  dorsal  filaments  arising  from  a  short  common  base,  the  other  two  being  separate 
for  their  entire  length;  these  filaments  as  long  as,  or  longer  than,  antennal  sheaths,  varying 
in  length  from  rather  short  to  a  longer  type.  Thoracic  dorsum  broad,  ample,  feebly  wrinkled 
transversely.  Leg  sheaths  (Plate  XX,  60)  with  all  the  tarsi  very  long  and  slender,  reaching 
almost  to  end  of  fifth  abdominal  segment.  Wing  sheaths  comparatively  narrow,  reaching 
base  of  third  abdominal  segment;  anal  angle  sharp;  venation  fairly  distinct. 

Abdomen  pale.  Intermediate  abdominal  segments  divided  into  two  annuli,  the  posterior 
ring  much  the  larger;  dorsa  of  segments  3  to  6  (Plate  XXII,  71),  and  sternum  of  segment 
6,  each  with  basal  annulus  tumid  and  with  two  transverse  rows  of  small  hooks  converging  at 
the  ends  to  inclose  a  linear  depressed  area;  these  areas  capable  of  contraction,  so  that  the 
hooks  of  each  row  are  united  or  approximated  with  those  of  the  opposite  row;  from  thirty 
to  thirty-five  hooks  in  each  row,  anterior  row  with  hooks  directed  backward,  posterior  row 
with  hooks  directed  forward;  on  seventh  segment,  only  the  anterior  row  of  hooks  present,  very 
slightly  arcuated,  the  lateral  hooks  smaller  than  those  near  middle  of  row;  caudad  of  this 


806  CHARLES  PAUL  ALEXANDER 

row  on  segment  7,  a  darkened,  transversely  rectangular  area  bearing  setiferous  punctures 
in  two  broken  rows,  the  posterior  row  the  more  complete;  posterior  rings  of  other  segments 
of  dorsum  bearing  setae  in  somewhat  the  same  arrangement.  Pleural  area  pale,  segments 
2  to  8  with  a  long,  delicate  seta  on  each  annulus.  Dorsum  of  segment  8  (Plate  XXII,  70)  with 
a  large  setiferous  tubercle  on  either  side,  this  tubercle  densely  covered  with  hairs  that  are 
longest  behind  and  shorter  in  front;  caudad  of  these  large,  blunt  knobs,  a  slender,  setiferous 
tubercle.  Last  segment  with  tergal  valves  chitinized,  elongate,  extreme  posterior  margin 
rounded  medially  and  feebly  bilobed,  lateral  angles  produced  caudad  and  dorsad  into  power- 
ful curved,  heavily  chitinized  hooks;  a  few  setae  at  about  midlength  of  these  hooks 
(Plate  XX,  62).  Sternal  valves  shorter,  slightly  bilobed  medially. 

Nepionotype. —  Ithaca,  New  York,  June  4,  1917. 

Neanotype. —  With  the  nepionotype. 

Paratypes  —  Topotypic,  May  1  to  June  10,  1917. 

Subtribe  Ellipteraria 

The  present  knowledge  of  the  immature  stages  of  the  genus  Elliptera 
is  due  entirely  to  the  work  of  Mik  (1886  b).  From  his  rather  detailed 
description  and  figures,  it  certainly  appears  that  the  group  should  receive 
coordinate  rank  with  Antocharia,  Limnobaria,  and  other  divisions  herein 
created.  The  genus  Elliptera  shows  peculiarities  of  structure  in  all  stages, 
but  many  features  of  its  organization  remind  one  forcibly  of  species  of 
Dicranomyia  (such  as  D.  simulans  and  D.  trinotata)  on  the  one  hand,  and 
of  Antocha  on  the  other;  and  it  may  be  that  the  genus  Elliptera  stands  in 
closer  relationship  to  Dicranomyia  than  is  now  believed. 

Genus  Elliptera  Schiner  (Gr.  I  omit,  or  ellipse  +  wing) 

1863     Elliptera  Schin.     Wien.  Ent.  Monatschr.,  vol.  7,  p.  222. 

1913    Ellipoptera  Bergr.    Ann.  Mag.  Nat.  Hist.,  8th  ser.,  vol.   11,  p.  576  (correct 
spelling). 

Elliptera  is  a  small  genus,  including  but  five  species  which  have  a  dis- 
continuous range,  two  being  found  in  Europe  and  three  in  western  North 
America.  The  commonest  of  the  North  American  species,  E.  clausa 
O.  S.,  was  found  on  wet  moss  in  the  spray  of  Vernal  Falls,  Yosemite  Valley, 
California  (Osten  Sacken,  1877:198).  The  only  information  available  on 
the  immature  stages  of  a  member  of  this  genus  is  that  furnished  by  Mik 
(1886b)  on  the  European  species  E.  omissa  Egg.  (quoted  subsequently 
by  Griinberg,  1910:31-32,  and  by  Malloch,  191 5-17 b:  226-227).  The 
description  and  account  as  given  below  are  based  entirely  on  Mik's 
paper. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  807 

Elliptera  omissa  Egg. 

1863    Elliptera  omissa  Egg.    Verh.  Zool.-Bot.  Ges.  Wien,  vol.  13,  p.  1108. 

Specimens  of  Elliptera  omissa  were  found  by  Mik  (1886b)  along  water- 
courses and  near  falls  in  mountainous  regions.  The  adult  flies  were 
noted  as  late  as  September  10,  swarming  about  the  waterfalls. 

Larvae  and  pupae  were  found  on  July  30  and  August  17  near  Salzburg, 
upper  Austria,  living  in  elongate  and  somewhat  depressed  cocoons  about 
10  millimeters  long  and  4  millimeters  broad  which  were  arranged  in  longi- 
tudinal rows  with  short  spaces  between.  These  cocoons,  which  were 
placed  with  the  current,  occurred  on  the  wet  walls  of  wooden  chutes  or 
runways  and  also  on  dripping  chalk  cliffs.  The  immature  stages  spend 
their  existence  in  these  small  cocoons  of  mud  and  silk.  When  ready  to 
emerge  as  an  adult,  the  pupa  makes  its  way  thru  the  end  of  the  cocoon 
away  from  the  current,  leaving  the  cast  skin  attached  to  the  opening.  The 
margins  of  the  large  pronotal  breathing  horns  of  the  pupa  are  finely  ser- 
rated and  are  presumably  used  in  making  this  opening  thru  the  cocoon. 
The  young  larva  probably  creeps  about  on  the  floor  of  the  runway,  feeding 
on  algae  growing  in  the  same  situation.  When  nearly  full-grown,  the 
larva  crawls  to  a  less  exposed  place  and  spins  its  cocoon.  Many  larvae 
and  pupae  are  killed  by  the  drying-out  of  their  haunts  when  the  water 
supply  becomes  insufficient  to  cover  them. 

Larva. —  Length,  7  mm. 

Diameter,  1.5  mm. 

Body  clearly  depressed  (Plate  XXIII,  72),  greenish  white,  scarcely  shiny,  with  delicate 
appressed  grayish  hairs  which  are  thicker  at  the  two  ends  of  the  body,  especially  on  last 
segment,  where  they  become  almost  villous.  Integument  very  transparent,  so  that  intes- 
tine  and  contents  show  thru,  the  intestine  narrowing  on  segment  6  and  thru  to  segment  8, 
where  it  broadens  out  and  almost  entirely  fills  the  ninth  and  tenth  segments.  On  sides  of 
prothorax  a  delicate,  long,  pale  hair;  on  remaining  segments  two  such  hairs.  On  each  of 
abdominal  segments  3  to  9,  on  dorsum  near  anterior  margin,  a  low  transverse  ridge  which 
is  thickly  set  with  short,  blackened  points;  on  sternum  of  each  of  same  segments,  a  similar 
welt  which  is  destitute  of  points.  In  male  larvae,  clawlike  appendages  of  genitalia  of  adults 
showing  thru  skin  on  ventral  side. 

Head  capsule  (Plate  XXIII,  73  and  74)  massive,  slightly  longer  than  broad,  black,  some- 
what shiny,  all  the  sclerites  compact  and  closely  united;  anterior  projecting  part  of  capsule  with 
margins  transparent,  rust-brown;  median  part  with  two  small  knobs,  laterad  of  which  are 
two  larger  projections  which  are  crowned  with  short  points;  capsule  weakly  keeled  behind 
on  dorsum  (Plate  XXIII,  74),  anterior  to  which  are  two  swollen  elevations;  on  hinder  mar- 
gin of  clypeus  a  styliform,  bristly  lobe,  easily  broken  off,  which  is  presumably  the  antenna. 


808  CHARLES  PAUL  ALEXANDER 

Labium  strongly  chitinized,  triangular,  split  longitudinally.  Mandible  (Plate  XXIII,  75) 
clawlike  in  appearance,  a  little  smaller  than  either  half  of  labium,  on  inner  face  with  a  chiti- 
nized projection  which  is  serrated.  Maxilla  indistinct,  the  palpi  coroniform.  Spiracular  disk 
(Plate  XXIII,  77)  with  four  lobes  whose  inner  faces  are  narrowly  lined  with  black  chitin; 
lobes  provided  with  lashes  of  long  gray  hairs;  dorsal  lobes  the  shorter  and  broader,  and  bear- 
ing on  their  inner  face  two  elongated  stigmata  which  are  margined  with  pale  rust-yellow. 

Pupa.' —  Length,  6.5  mm. 

Pronotal  breathing  horns  (Plate  XXIII,  76)  large,  ear-shaped;  bright  yellowish  white  in 
color,  in  contrast  to  dirty  yellowish  brown  skin  of  head,  thorax,  and  appendages;  each  horn 
consisting  of  two  parts:  the  dorsal  side,  appearing  smooth  and  homogeneous;  and  the  ventral 
side,  with  two  longitudinal  furrows  converging  toward  apices,  and  with  abundant  elongate 
tubercles,  under  low  magnification  this  part  appearing  pitted  because  of  the  spaces  between 
these  tubercles.  Margin  of  breathing  horn  chitinized  and  very  finely  notched.  On  outer 
basal  part  of  each  ear  a  parchment-like  lobe,  which  joins  ear  to  side  of  prothorax;  in  addition 
to  this,  each  ear  at  base  is  drawn  out  into  an  almost  rectangular  lobe  which  is  closely  approxi- 
mated to  pronotum.  Leg  sheaths  extending  about  to  base  of  fifth  abdominal  segment. 
Abdomen  distinctly  depressed,  greenish  white  in  color;  segments  3  to  7  on  both  dorsum  and 
sternum  near  base  with  a  double  cross-row  of  spicules  which  present  a  comblike  appearance 
(Plate  XXIII,  78),  those  on  dorsal  segments  being  somewhat  stronger.  Female  pupa  with 
acidothecae  grown  together  on  inner  face,  at  outer  angle  of  each  a  chitinized  hook  which  is 
curved  upward.  Male  pupa  with  ventral  side  of  last  segment  produced  into  two  chitinized 
points  which  are  bent  toward  each  other  and  almost  touch,  these  inclosing  clasping  organs 
of  adult  male  and  hooks  of  male  larva  as  described  above. 

Subtribe  Limnobaria 

The  subtribe  Limnobaria  includes  about  ten  genera,  which  are  very 
closely  related  to  one  another  and  whose  limits  are  as  yet  not  clearly 
defined.  Many  of  the  generic  distinctions  are  based  on  male  characters 
of  wing  form,  venation,  or  antennal  structure.  The  genera  are  often  very 
large  and  it  is  difficult  to  give  satisfactory  characters  to  separate  their 
immature  stages.  The  keys  to  the  genera,  given  below,  will  unquestion- 
ably need  much  revising  when  a  larger  number  of  forms  are  studied. 

Larvae 

1.  Form  stout;  teeth  of  mandibles  and  of  mentum  usually  more  numerous;  mentum  more 

pointed  anteriorly Limnobia  Meig.  (p.  809) 

Form  usually  more  slender;  teeth  of  mandibles  and  of  mentum  usually  fewer;  mentum 
transverse  or  subtransverse 2 

2.  Mandibles  very  broad,  flattened,  with  three  ventral  cutting  teeth;  mentum  about  trans- 

verse, with  nine  or  eleven  teeth Rhipidia  Meig.  (p.  825) 

Mandibles  more  slender,  with  usually  four  or  five  cutting  teeth;  mentum  usually  a  little 
pointed  anteriorly,  with  about  eleven  teeth Dicranomyia  Steph.  (p.  819) 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  809 

Pupae 

1.  The  five  basal  abdominal  segments  on  both  dorsum  and  venter  with  a  comb  of  small, 

blunt  teeth;  wing  sheaths  showing  an  ocellate  pattern;  pupae  living  beneath  bark  of 

coniferous  trees Discobola  O.  S.  (p.  815) 

Abdominal  segments  provided  with  basal  transverse  welts  of  microscopic  points  on  seg- 
ments 3  to  7 ;  wing  pattern  not  ocellate 2 

2.  Pronotal  breathing  horns  long  and  narrow,  about  three  times  as  long  as  broad. 

Rhipidia  Meig.  (p.  825) 
Pronotal  breathing  horns  short  and  broad,  length  and  breadth  not  greatly  different 3 

3.  Size  large  (usually  over  10  mm.) ;  breathing  horns  often  broader  than  long. 

Limnobia  Meig.  (p.  809) 
Size  smaller  (usually  under  10  mm.) ;  breathing  horns  usually  as  long  as  broad. 

Dicranomyia  Steph.  (p.  819) 

Genus  Limnobia  Meigen  (Gr.  swamp  +  I  live) 

1800    Amphinome  Meig.    Nouv.  Class.  Mouch.,  p.  15  (nomen  nudum,  preoccupied  in 

Annelida). 

1803    Limonia  Meig.     Illiger's  Mag.,  vol.  2,  p.  262. 
1818    Limnobia  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  1,  p.  116. 
1818     Unormjia  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  1,  p.  116. 
1856    Limnomyza  Rond.     Dipt.  Ital.  Prodr.,  vol.  1,  p.  185. 

Larva." —  Form  stout,  terete.  Abdominal  segments  with  dorsal  and  ventra*  transverse 
welts  covered  with  chitinized  points  on  basal  rings.  Spiracular  disk  surrounded  by  indis- 
tinct lobes,  the  spiracles  large.  Head  capsule  large,  massive,  the  dorsal  plate  narrowed 
behind  and  more  or  less  bifid  at  its  tip.  Labrum  transversely  oval,  with  sensory  bristles 
near  margin.  Mandible  blunt,  with  from  four  to  seven  cutting  teeth.  Maxilla  simple, 
cardo  and  stipes  large.  Antenna  with  apical  papilla  button-like.  Mentum  broad,  margin 
with  from  eleven  to  thirteen  teeth.  Hypopharynx  a  circlet  of  two  chitinized  plates,  each 
with  from  twelve  to  fifteen  teeth. 

Pupa.- —  No  cephalic  crest.  Pronotal  breathing  horns  flattened,  earlike,  as  broad  as,  or 
broader  than,  long.  Meson otum  unarmed.  Abdomen  with  transverse  welts  of  fine  hooks 
on  basal  annuli  of  tergites  3  to  7  and  stern ites  5  to  7.  Two  tiny  ^piracies  on  dorsum  of 
eighth  abdomina)  segment. 

Limnobia  is  a  rather  small  genus  (comprising  about  forty-five  species) 
of  usually  large  and  handsome  flies.  The  species  are  most  numerous 
thruout  the  Holarctic  and  Ethiopian  regions.  The  immature  stages  have 
a  wide  range  of  habitat. 

Of  the  European  species,  Limnobia  bifasciata  Schr.  [=  L.  xanthoptera 
Meig.]  is  characteristically  fungicolous,  the  larvae  occurring  in  various 
species  of  Agaricus  and  related  genera  as  stated  by  Stannius,  Pastejrik, 
De  Meijere,  and  other  investigators.  L.  decemmaculata  Lw.  occurs  in 
fungi  (Daedalea  and  similar  species),  as  recorded  by  Loew  (1873)  and  by 
Verrall  (1912).  L.  quadrimaculata  (Linn.)  [=  L.  annulus  Meig.]  often 
occurs  in  tree  fungi  but  is  not  confined  to  this  habitat.  This  species, 


810  CHARLES  PAUL  ALEXANDER 

L.  macrostigma  Schum.,  and  L.  obscuricornis  Bel.  are  often  found  in  decay- 
ing, principally  deciduous,  wood.  L.  tripunctata  Fabr.,  L.  sexpunctata 
Fabr.  [=L.  nigropunctata  Schum.],  L.  flavipes  Fabr.,  and  L.  nubeculosa 
Meig.  are  found  in  humous  earth  and  beneath  leaves  in  woods.  The 
pupal  duration  of  L.  quadrimaculata  is  from  eight  to  twelve  days,  and  this 
species,  as  well  as  others  of  the  genus,  pupates  in  the  ground,  inclosed  in 
delicate  silken  cases  which  are  covered  with  particles  of  earth  and  other 
matter. 

In  America,  L.  triocellata  0.  S.  is  characteristically  fungicolous.  L. 
cinctipes  Say,  and  presumably  L.  immatura  0.  S.,  are  found  both  in  fungi 
and  in  decaying  wood.  L.  indigena  0.  S.  has  been  found  in  living  tulip 
roots  from  Greenville,  South  Carolina  (Greene,  ms.).  L.  fallax  Johns., 
and  presumably  L.  solitaria  O.  S.,  live  in  organic  mud  near  water.  L. 
parietina  0.  S.  probably  has  a  similar  habitat,  since  it  was  found  in  tent 
traps  set  over  Beaver  Meadow  Brook  in  the  Adirondacks  (Needham, 
1908  a:  171). 

Limnobia  cinctipes  Say 

1823    Limnobia  cinctipes  Say.    Journ.  Acad.  Nat.  Sci.  Phila.,  vol.  3,  p.  21,  no.  4. 

Limnobia  cinctipes  is  one  of  the  largest  and  commonest  American  species 
of  the  genus  Limnobia.  It  has  been  reared  many  times,  some  of  the 
records  being  as  follows: 

On  July  23,  1883,  Pergande  collected  larvae  in  an  old  fungus  growing 
on  rotten  wood.  On  July  20,  1886,  the  same  species  of  larvae  was  found 
constructing  silken  cases  thru  the  fungus,  and  later  in  the  ground  for 
pupation.  The  pupae  were  active,  and  were  able  to  draw  back  and  forth 
in  their  tubes.  Adults  began  to  issue  on  July  28,  showing  the  pupal 
stage  in  this  case  to  be  not  more  than  eight  clays. 

On  April  25,  1912,  a  number  of  specimens  of  this  species  were  received 
from  W.  H.  Shideler,  of  Miami  University,  Ohio.  The  specimens  were 
taken  at  Oxford,  Ohio,  on  April  20,  when  several  hundred  larvae  and 
pupae  were  found  in  an  old  dry  log.  The  young  pupae  in  the  wood  are 
not  covered  with  particles  of  debris,  but  the  older  pupae  are  inclosed  in 
a  case  which  is  covered  with  wood  fragments,  only  the  top  of  the  head  and 
the  tip  of  the  abdomen  projecting  beyond  the  case.  When  the  pupae 
are  about  to  transform,  the  insect  emerges  to  about  half  its  length  and  the 


THE  CRANE-FLIES  OF  NEW  YORK — PART  II  811 

skin  splits  down  the  dorsum,  the  pupal  skin  remaining  in  place  after  the 
adults  have  emerged. 

On  September  15,  1912,  many  full-grown  larvae  were  found  in  a  fleshy 
species  of  Fomes  near  Glovers ville,  New  York,  where  they  were  asso- 
ciated with  a  much  larger  number  of  larvae  of  Ula  elegans  and  a  much 
lesser  number  of  Limnobia  triocellata.  When  about  to  pupate,  the  larva 
becomes  pale  green  in  color  and  incases  itself  completely  in  a  silken  sheath 
which  is  covered  with  particles  of  sand  and  other  debris.  As  the  pupa 
grows  older,  the  case  becomes  harder  and  more  rigid.  Numerous  little 
mites  are  to  be  found  running  up  and  down  over  these  pupae,  more 
especially  at  the  head  end,  and  possibly  seeking  ingress  into  the  insect. 
One  young  pupa  had  a  piece  of  cloth  adhering  to  the  side  of  its  case.  The 
pupal  stage  lasts  about  five  days. 

Larva. —  Length,  18-22  mm. 

Diameter,  2.5-3.2  mm. 

Coloration  light  yellow  to  greenish,  the  setiferous  transverse  welts  at  base  of  abdominal 
segments  brownish. 

Body  terete,  abdominal  segments  subdivided  into  two  narrow  basal  rings  and  a  broad 
posterior  ring.  Abdominal  segments  1  to  7  with  a  broad  basal  welt  on  tergites  and  sternites, 
that  of  the  first  segment  much  smaller;  these  welts  densely  covered  with  microscopic  hooks; 
on  the  last  two  thoracic  segments,  welts  indicated  by  very  narrow  lines.  Cauda  blunt, 
obliquely  truncated.  Spiracular  disk  (Plate  XXIV,  83,  and  Plate  XXV,  93)  surrounded 
by  indistinct  lobes,  the  ventral  margin  projecting  far  caudad  and  indistinctly  divided  into" 
two  short  lobes;  lateral  lobes  very  blunt;  dorsal  lobes  short  and  blunt,  often  divided  into 
two  smaller  lobes.  Spiracles  oblong  or  elliptical,  placed  obliquely.  Gills  four,  blunt  and 
rounded,  formed  for  propulsion  rather  than  for  respiration. 

-  Head  capsule  (Plate  XXIV,  79)  very  much  as  in  Antocha,  the  dorsal  plate  narrowed  behind 
and  somewhat  bifid  at  apex;  lateral  plates  shaped  like  a  mussel  shell,  curved  around  to  form 
mentum.  Labrum  (Plate  XXV,  86)  distinct,  oval,  the  anterior  margin  fringed  with  delicate 
hairs,  the  hairs  at  the  lateral  margins  longer  and  coarser;  on  either  side  near  anterior  margin, 
a  blunt  tubercle  with  three  sensory  bristles;  just  laterad  of  this  a  stout  seta;  along  anterior 
margin,  four  sensory  setae  which  are  subequally  spaced.  Epipharynx  densely  hairy. 
Clypeus  broader  than  labrum,  with  a  seta  at  each  outer  anterior  angle  and  two  more  on 
either  side  near  posterior  margin.  Mentum  (Plate  XXIV,  81)  elongate-triangular,  not  com- 
pletely divided  into  halves  but  deeply  split  behind,  with  an  outer  plate  running  cephalad  into 
a  long,  broad  point;  behind  this  another  plate  with  the  margins  toothed,  there  being  about 
five  or  six  long,  acute  teeth  on  either  side.  Hypopharynx  (Plate  XXIV,  80)  with  two 
rows  of  teeth  forming  a  circlet,  into  which  duct  of  salivary  gland  opens;  anterior  row  having 
about  nine  large,  blunt  teeth,  with  about  six  smaller  teeth  on  either  side,  these  latter  sharper- 
pointed  and  more  crowded;  posterior  row  having  long,  pointed  teeth,  about  twelve  in 
number.  Antenna  (Plate  XXIV,  82)  two-segmented,  the  basal  segment  chitinized,  elongate- 


812  CHARLES  PAUL  ALEXANDER 

cylindrical,  the  second  segment  flattened,  shaped  somewhat  like  a  door  knob;  a  few  sensory 
projections.  Mandible  (Plate  XXV,  87  and  88)  powerful,  produced  into  a  strong  apical 
point,  with  about  four  or  five  blunt  or  irregular  inner  teeth  and  a  strong  dorsal  tooth  on 
outer  margin  before  tip.  Maxilla  (Plate  XXV,  87)  large,  the  outer  margin  thickened,  sub- 
chitinized;  palpi  at  apex  small,  shaped  like  half  a  pill  box,  with  a  few  sensory  papillae 
at  tip. 

Pupa. —  Length,  18-20  mm. 

Width,  d.-s.,  2.8-3  mm. 
Depth,  d.-v.,  3.1-3.3  mm. 

Entire  head  and  thorax,  including  leg  and  wing  sheaths,  light  brown,  the  thoracic  dorsum 
somewhat  darker-colored,  the  wings  more  yellowish  brown;  abdomen  pale  light  green,  the 
segments  with  the  submedian  brown  band  interrupted  on  pleural  region;  tip  of  abdomen 
brownish,  chitinized. 

Form  stout  (Plate  XXV,  89);  body  destitute  of  noticeable  setae. 

Head  flattened  (Plate  XXIV,  84) .  Cephalic  crest  lacking;  forehead  with  a  shallow  V-shaped 
notch  between  antennal  bases.  Eyes  of  male  large,  the  front  narrowed,  with  points  of 
tentorium  close  to  inner  margin  of  eye;  eyes  of  female  more  widely  separated.  Antenna 
rather  short,  ending  just  beyond  wing  root.  Labrum  short,  obtuse.  Labial  lobes  contigu- 
ous, divergent,  blunt  at  their  tips,  posterior  margin  a  little  convex  medially.  Lobes  of 
maxillary  palpi  large,  subquadrate.  Cheek  with  a  large,  flattened  ledge  overlying  joint  of 
fore  legs. 

Thorax  very  gibbous.  A  distinct  anterior  median  carina  between  breathing  horns. 
Pronotal  breathing  horns  (Plate  XXV,  90)  flattened,  earlike,  broader  than  long,  directed 
slightly  proximad,  margin  with  a  row  of  breathing  tubercles,  outer  face  wrinkled.  Wing 
sheaths  reaching  base  of  third  abdominal  segment.  Leg  sheaths  reaching  base  of  fourth 
abdominal  segment  or  a  little  longer;  tarsi  ending  about  on  a  level,  or  sloping  gradually  from 
short  hind  tarsi  to  long  fore  tarsi. 

Abdominal  segments  indistinctly  subdivided  into  three  rings;  on  tergites  3  to  7,  and 
sternites  5  to  7,  basal  ring  with  a  transverse  welt  which  is  densely  covered  with  short  hairs 
or  hooks,  these  welts  tapering  gradually  to  ends;  sternites  of  segments  3  and  4  having  incom- 
plete welts  on  either  side  of  leg  sheaths;  band  on  tergum  of  segment  7  not  broken  medially, 
but  a  little  constricted  in  some  specimens;  in  older  pupae  the  other  annuli,  especially  the 
posterior  one,  variously  darkened  on  dorsum  and  venter.  Female  cauda  (Plate  XXIV,  85) 
with  the  acidothecae  short,  the  sternal  valves  the  shortest,  the  tergal  valves  a  little  longer; 
prominent  lateral  lobes  at  base  of  tergal  valves,  and  a  slightly  smaller  but  very  broad  one 
on  each  side  of  tergal  valves  at  about  midlength.  Male  cauda  (Plate  XXV,  91)  similar  to 
that  of  female,  but  the  dorsal  lobes  (Plate  XXV,  92)  much  shorter,  not  longer  than  the  ven- 
tral lobes,  and  separated  by  a  U-shaped  notch;  ventral  lobes  approximated,  each  ending 
in  a  small,  blunt  tubercle.  Two  small  circular  spiracles  on  dorsum  of  segment  8,  these  a 
little  more  widely  separated  in  male  than  in  female. 

Nepionotype. —  Gloversville,  New  York,  October  26,  1912. 
Neanotype. —  Female  pupa  with  type  larva. 

Paratypes. —  Several  larvae  and  pupae  with  types  and  from  Oxford,  Ohio,  April  20, 
1912. 


THE  CEANE-FLIES  OF  NEW  YORK  —  PART  II  813 

Limnobia  fallax  Johns. 

1909    Limnobia  fallax  Johns.    Proc.  Boston  Soc.  Nat.  Hist.,  vol.  34,  p.  125. 

Limnobia  fallax  belongs  to  the  solitaria  group  and  is  apparently  more 
Austral  in  its  distribution  than  the  other  members  of  this  group  (L.  soli- 
taria O.  S.,  L.  hudsonica  O.  S.). 

Larvae  and  pupae  were  found  by  Dr.  Johannsen  near  Ithaca,  New  York, 
July  20-26,  1905.  They  were  wrapped  in  silken  cases  covered  with  earthy 
matter,  and  were  removed  from  the  soil  near  a  brook. 

Larva. —  Length,  contracted,  8-8.2  mm. 
Diameter,  1.2-1.3  mm. 

Coloration  white. 

Form  stout  and  short,  body  terete.  Transverse  welts  with  chitinized  points  on  abdominal 
segments  2  to  7,  those  on  dorsal  surface  broad,  those  on  ventral  surface  narrower.  Spiracular 
disk  blunt,  surrounded  by  four  indistinct  lobes,  the  lateral  pair  the  largest  and  capable  of 
close  approximation,  closing  the  large  yellow  spiracles.  Anal  gills  indistinct. 

Head  capsule  of  usual  Limnobia  type.  Labrum  (Plate  XXVI,  95)  broadly  oval,  lateral 
angles  and  disk  of  epipharynx  with  tufts  of  long  hairs;  anterior  margin  fringed  with  short 
setae;  about  eight  sensory  bristles  and  papillae  along  anterior  margin.  Mentum  (Plate 
XXVI,  96)  large,  triangular,  running  out  into  a  long  median  apical  point;  lateral  margins 
with  about  six  or  seven  flattened  subacute  teeth  on  each  side.  Hypopharynx  as  in  this  group: 
a  collar  formed  of  two  chitinized,  comblike  plates;  the  first  plate  rectangular,  its  face  covered 
with  flattened  scales,  anterior  margin  with  about  fourteen  acute  pointed  teeth,  the  two 
outermost  much  the  smaller,  acicular;  the  second  plate  a  narrow  band  of  chitin  similarly 
ioothed,  the  teeth  at  each  end  large,  flattened,  the  next  tooth  very  narrow,  acicular,  the 
remaining  teeth,  ten  in  number,  flattened,  acute,  the  middle  teeth  a  little  shorter  and  broader. 
Antenna  (Plate  XXVI,  97)  with  basal  segment  elongate,  cylindrical,  and  apical  papilla  or  seg- 
ment very  tiny,  disklike.  Mandible  (Plate  XXVI,  98)  large,  moderately  broad,  with  apical 
tooth  prominent,  two  large  dorsal  teeth,  and  a  row  of  about  five  or  six  comblike  teeth  along 
ventral  cutting  edge,  the  most  basal  being  short  and  blunt.  Maxilla  (Plate  XXVI,  99) 
about  as  in  this  tribe,  cardines  and  stipites  large  and  simple;  palpus  large. 

Pupa. —  Length  of  cast  pupal  skin,  10-13  mm. 

Labrum  (Plate  XXVI,  100)  triangular,  apex  obtuse.  Labial  lobes  prominent,  subquadrate. 
Posterior  margin  convex  medially.  Pronotal  breathing  horns  (Plate  XXVI,  101)  flattened, 
subcircular  in  outline,  with  an  outer  marginal  row  of  breathing  tubercles.  Leg  sheaths 
ending  just  before  apex  of  fourth  abdominal  segment;  tips  of  tarsi  about  on  a  level,  or  those 
of  fore  legs  a  little  the  longer.  Band  of  spicules  on  seventh  tergite,  slightly  constricted 
medially.  Female  cauda  (Plate  XXVI,  102  and  103)  with  tergal  valves  a  little  longer  than 
sternal  valves,  and  more  acute  at  their  tips;  a  small  tubercle  on  outer  margin  of  outer  lobes 
Defore  tips.  „ 

Nepionotype.—  Ithaca,  New  York,  July  21,  1905. 
Neanotype. —  Cast  pupal  skin  with  type  larva. 
Paratypes. —  One  larva  and  three  pupal  skins. 


814  CHARLES  PAUL  ALEXANDER 

Limnobia  triocellata  0.  S. 

1859    Limnobia  triocellata  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  216. 

Limnobia  triocellata  is  a  common  crane-fly  in  eastern  North  America. 
It  is  closely  allied  to  the  European  L.  bifasciata  Schr.,  the  immature  stages 
of  which  have  long  been  known. 

Johnson  (1906:2)  found  larvae  of  this  species  in  a  fungus  at  Riverside, 
Massachusetts,  on  August  21,  1904,  which  pupated  on  the  22d  and  emerged 
on  the  30th  and  31st,  thus  giving  a  pupal  duration  of  about  nine  days. 
Malloch  (1915-17  b :  215-216)  found  larvae  and  cast  pupal  skins  in  an  Agari- 
cus  at  Urbana,  Illinois,  in  September  of  1915.  The  writer  found  larvae 
of  Limnobia  triocellata  in  a  species  of  Fomes,  associated  with  the  larvae 
of  L.  cinctipes  and  Ula  elegans,  at  Gloversville,  New  York,  on  September 

15,  1912.     C.  H.  Popenoe  found  larvae  at  Great  Falls,  Virginia,  on  Sep- 
tember 8,  1912,  in  the  fungi  Hypomyces  Lactifluorum   (Schw.)  Tul.  and 
Armillaria  sp.,  the  adult  flies  emerging  on  October  7,  1912.     Other  speci- 
mens from  the  same  place  found  on  October  9,  1913,  in  a  species  of  Clito- 
cybe,  produced  adults  on  October  20.     Scores  of  specimens  were  taken  in 
Boletus  felleus  at  Bradley  Hill,   Maryland,  the  flies  emerging  on  July 

16,  1914. 

Larva. —  Length,  10-18  mm. 

Diameter,  1.2-1.4  mm. 

Coloration  a  little  more  yellowish  than  that  of  Limnobia  fallax.  Species  very  close  to 
fallax  in  all  details.  Ventral  welt  on  abdominal  segment  1  well  developed,  but  dorsal  welt 
lacking  or  very  reduced.  Spiracular  disk  (Plate  XXVI,  94)  rather  large,  exposed;  circular 
spiracles  large,  separated  by  a  distance  less  than  the  diameter  of  one;  lobes  surrounding 
disk  small  and  indistinct. 

Pupa. —  Length,  12-15  mm.  Not  very  different  from  other  species  of  genus  described 
herein. 

Nepionotype. —  Great  Falls,  Virginia,  September  28,  1913. 
Paratypes. —  With  the  type. 

Genus  Libnotes  Westwood  (derivation  obscure) 
F  1876    Libnotes  Westw.    Trans.  Ent.  Soc.  Lond.,  p.  505. 

Libnotes  is  a  small  genus  (about  thirty-five  species)  of  rather  large 
crane-flies,  which  are  chiefly  Oriental  in  their  distribution  altho  three 
species  occur  in  South  and  Central  Africa.  The  species  Libnotes  per- 
kinsi  (Grimsh.)  has  been  considered  as  being  a  Limnobia,  but  it  seems 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  815 

to  the  writer  that  the  present  generic  reference  is  more  nearly  correct, 
altho  the  distinctions  between  Limnobia  and  some  species  of  Libnotes 
are  very  poorly  marked.  L.  perkinsi  was  bred  from  larvae  in  damp  moss 
(Perkins,  1913  :clxxxii,  as  Limnobia),  and  in  a  letter  to  the  writer  Mr.  O.H. 
Swezey  states  that  he  has  reared  this  species  from  larvae  in  decaying 
vegetation  and  in  the  accumulation  of  debris  behind  old  leaf -sheaths  on 
banana  plants  (Musa,  Scitamineae)  in  the  Hawaiian  Islands. 

Genus  Discobola  Osten  Sacken  (Gr.  discus  +  I  throw) 

1865     Discobola  0.  S.     Proc.  Ent.  Soc.  Phila.,  p.  226. 

1869     Trochobola  O.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  98. 

Discobola  is  a  well-marked  genus  including  about  eight  described 
species,  which  are  most  numerous  in  the  Australasian  region,  two  species 
only  being  found  in  Europe  and  two  others  in  America.  The  adults  of 
the  commoner  American  species,  D.  argus  (Say),  are  not  rare.  They  are 
most  numerous  in  late  summer,  and  are  often  found  resting  on  the  stumps 
and  trunks  of  coniferous  trees,  especially  white  pine  (Pinus  Strobus 
Linn.). 

The  immature  stages  of  D.  caesarea  (O.  S.)  were  found  by  Mik  (1884) 
in  Austria,  living  in  decaying  pine  stumps  from  which  the  bark  had  been 
removed.  The  following  account  is  taken  entirely  from  Mik's  paper: 

Male  pupa. —  Body  cylindrical,  slender  (9.2  mm.  long,  1.5  mm.  in  diameter).  Head, 
prothorax,  mesothorax,  leg  sheaths,  and  wing  sheaths  chitinized,  dark  brown,  shiny,  the 
last-named  somewhat  brighter  than  the  others,  the  leg  sheaths  somewhat  darker  at  their 
'tips.  Eyes  kidney-shaped,  strongly  shiny,  blackish,  between  them  a  small,  triangular, 
blackish  brown  spot.  Prothoracic  breathing  horns  dull-colored,  dark  rust-brown  at  base, 
becoming  a  brighter  rust-brown  more  distally,  compressed  laterally,  tuberculate,  with  margin 
indented.  Prothorax  carinate,  rust-yellow,  margined  on  both  sides  by  dull  reddish  brown 
tubercles.  In  fully  colored  specimens,  forehead  and  leg  sheaths  blackish  brown,  wing  pat- 
tern indicated  on  sheaths  as  somewhat  diffused  rings.  Leg  sheaths  reaching  end  of  abdominal 
segment  3,  wing  sheaths  reaching  end  of  abdominal  segment  1.  Metathorax  and  abdomen 
thin-skinned,  the  former  verdigris-colored,  the  latter  white  or  somewhat  yellowish  green; 
metathorax  resembling  an  abdominal  segment,  but  its  posterior  margin  is  unarmed,  while 
the  first  to  the  fifth  abdominal  segments  on  both  dorsum  and  venter  bear  a  comb  of  very  small, 
short,  blunt  teeth,  which  are  closely  approximated;  these  teeth  chitinized  and  rusty  brown  at 
their  tips,  giving  to  abdomen  the  appearance  of  having  brown  incisions;  these  transverse 
rows  of  teeth  interrupted  at  pleura.  Sixth  abdominal  segment  pale  thruout  and  lacking 
the  comb.  Seventh  segment  shorter  and  narrowed  on  dorsum,  pale,  bearing  on  sternum 
a  rust-yellow  chitinized  plate  which  is  narrowed  anteriorly,  leaving  an  uncolored  triangular 
area  on  either  side  at  base  of  segment.  Eighth  segment  swollen  to  include  genitalia,  the 


816  CHARLES  PAUL  ALEXANDER 

two  basal  parts  ellipsoidal,  strongly  shiny,  rust-yellow,  somewhat  darker  at  tips,  the  apical 
parts  small  and  knotlike,  bluntly  rounded;  segment  bearing  on  dorsum  a  weak  triangular 
piece  at  its  base;  between  apical  parts  of  genitalia  are  inserted  two  small  chitinized  shields; 
on  venter,  between  basal  parts,  sheath  of  penis  is  inserted. 

Female  pupa.- —  Body  resembling  that  of  male,  but  longer  and  somewhat  stouter  (length 
10.5  mm.,  diameter  1.8  mm).  Leg  sheaths  extending  to  just  beyond  midlength  of  abdominal 
segment  2.  Seventh  abdominal  segment  shortened  and  somewhat  narrowed,  on  dorsum 
largely  pale,  with  a  narrow  chitinized  margin  only  on  lateral  parts,  so  that  the  unchitinized 
part  forms  a  triangle  with  the  apex  directed  backward;  on  sternum  this  segment  almost 
completely  chitinized,  rust-yellow,  only  a  small  triangular  area  at  base  on  either  side  remain- 
ing uncolored;  chitinized  plate  separated  from  plate  of  next  segment  only  by  an  incomplete 
segmentation,  swollen,  and  bearing  two  longitudinal  impressions.  Eighth  segment  bearing 
on  its  dorsal  surface  the  dorsal  valves  of  ovipositor,  fused  at  their  base,  chitinized  thruout, 
rust-yellow  in  color;  segment  bearing  on  its  ventral  surface  a  depressed  conical  chitinized 
plate  of  a  rust-yellow  color,  and  with  transverse  impressed  wrinkles;  on  either  side  a  small, 
dark,  chitinized,  lower  valve  of  ovipositor.  Other  characters  as  in  male.  (When  the  pupae 
are  placed  in  alcohol,  the  green  of  the  metathorax  and  the  abdomen  disappears  and  is 
replaced  by  a  yellowish  white  color.) 

Pupae  were  collected  in  large  numbers  in  a  pine  wood  near  Hammern 
in  Freistadt  (upper  Austria)  in  the  latter  days  of  August,  1882.  The 
pupae  live  in  pine  stumps,  near  the  ground,  where  the  bark  has  been 
removed,  more  especially  in  situations  where  the  wood  is  somewhat  sappy 
and  not  yet  completely  decayed.  Those  found  were  not  deep  in  the  wood. 
Their  presence  was  discovered  by  finding  the  teneral  adults  on  and  near 
a  stump,  and  many  cast  skins  of  the  pupae  projecting  horizontally,  the 
caudal  end  of  the  body,  up  to  the  leg  sheaths,  adhering  to  the  wood.  No 
emergence  holes  were  found  on  the  cut  surface  of  the  stump.  The  adults 
at  first  have  a  very  long,  pale  abdomen,  which  is  of  a  verdigris  color, 
most  intensive  at  the  base  and  paler  toward  the  tip.  The  pupae  that 
were  found  transformed  as  adults  in  from  one  to  three  days. 

Genus  Qeranomyia  Haliday  (Gr.  crane  +  fly) 

1833  Gerarwmyia  Hal.     Ent.  Mag.,  vol.  1,  p.  154. 

1835  Limnobiorhynchus  Westw.     Ann.  Soc.  Ent.  France,  vol.  4,  p.  683  (spurious  name). 

1838  Aporosa  Macq.     Dipt.  Exot.,  vol.  1,  part  1,  p.  62. 

1865  Plettusa  Phil.    Verh.  Zool.-Bot.  Ges.  Wien,  vol.  15,  p.  597. 

Geranomyia  is  a  rather  extensive  genus  including  about  eighty  species, 
which  are  most  abundant  in  the  tropics  of  America,  Asia,  and  Australia. 
On  the  African  continent  the  genus  is  apparently  less  common.  The 
adult  flies  have  an  elongate  rostrum  which  is  used  for  sucking  nectar 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  817 

from  tubular  flowers.  The  various  species  of  the  genus  have  been  recorded 
as  feeding  on  a  wide  range  of  plant  species,  which  have  been  indicated  by 
Knab  (1910)  and  by  Alexander  (1916  b:  486^493)  and  may  be  summarized 
as  follows: 

Species  Plants  frequented 

Geranomyia  canadensis  Compositae  —  Eupatorium,   Solidago,  Aster, 

Silphium,   Rudbeckia,  Verbesina,   Cacalia, 
and  similar  species 
Geranomyia  diversa  Compositae  —  Solidago,  Erigeron 

Umbelliferae  —  Daucus 

Geranomyia  virescens  Lauraceae  —  Persea 

Geranomyia  rostrata  Compositae  —  Eupatorium,    Solidago,    Heli- 

anthus 

For  many  years  nothing  was  known  concerning  the  immature  stages 
of  any  species  of  Geranomyia.  In  1917,  J.  R.  Malloch  found  larvae  and 
pupae  of  G.  canadensis  at  Urbana,  Illinois.  Mr.  Malloch  and  the  writer 
have  in  press  a  detailed  paper  on  the  immature  stages  of  this  species,  and 
the  following  brief  account  is  abstracted  from  this  paper  and  included 
herewith  in  order  to  complete  the  data. 

Mr.  Malloch  found  the  larvae  on  the  grounds  of  the  Floriculture  Depart- 
ment of  the  University  of  Illinois.  There  is  a  small  bubbling  fountain 
here,  the  waste  water  from  which  flows  along  an  open  gutter.  In  this 
gutter  the  immature  stages  of  G.  canadensis  lived  among  the-  vegetable 
growth  and  diatomaceous  ooze  in  the  bottom  of  the  trough.  Mr.  Malloch 
and  the  writer  found  this  same  species  in  Union  County,  southern 
Illinois,  in  1919.  Here  larvae  and  pupae  occurred  on  the  face  of  rocks 
where  the  surface  was  continually  damp  with  percolating  water.  A 
railroad  bank  had  been  formed  by  piling  up  slabs  of  limestone  to  a  height 
of  about  four  feet.  In  the  irregularities  and  crevices  of  these  pieces 
of  limestone,  the  larvae  of  Geranomyia  were  living  in  delicate  silken 
tubes  covered  with  a  deposit  of  silt  and  diatoms.  They  emerged  from 
their  cases  to  feed  on  the  exposed  surface  of  the  wet  rocks  during  twilight, 
and  even  during  the  hours  of  sunlight,  but  upon  being  disturbed  or  alarmed 
they  retreated  with  great  agility  into  their  tubes.  The  pupae  are  found 
in  short,  nearly  vertical  burrows  in  the  same  situations  as  the  larvae; 
here  they  rest  with  only  the  long,  conspicuous  breathing  horns  projecting 
from  the  entrance  to  the  burrow.  When  transformation  takes  place, 
the  pupal  skin  projects  from  the  mouth  of  the  burrow  nearly  to  the  ends 


818  CHARLES  PAUL  ALEXANDER 

of  the  wing  sheaths.  The  number  of  larvae  vastly  exceeds  the  number 
of  pupae,  and  this  would  seem  to  indicate  that  the  pupal  existence  is  of 
very  short  duration,  else  this  stage  would  be  found  oftener. 

Larva. —  Length,  12-12.5  mm. 
Diameter,  0.8-0.9  mm. 

Coloration  grayish  subhyaline;  a  large  orange  area  on  posterior  lateral  parts  of  prothorax; 
abdominal  welts  dark  brown. 

Form  moderately  long  and  slender;  thoracic  segments  gradually  decreasing  in  length  from 
prothorax  to  mesothorax;  abdominal  segments  gradually  elongated  to  the  fifth,  thence  short- 
ened to  end  of  abdomen.  Ventral  surface  of  meso-  and  metathorax  and  of  first  eight  abdom- 
inal segments  provided  with  a  basal  transverse  welt  which  is  densely  set  with  microscopic 
points;  on  dorsal  surface  these  bands  smaller,  occurring  on  metathorax  and  on  abdominal 
segments  2  to  8,  not  connected  with  sternal  bands  except  on  metathorax  and  on  eighth 
abdominal  segment.  Spiracular  disk  similar  to  that  in  Dicranomyia;  the  usual  ventral 
lobes  represented  only  by  two  small,  dusky,  setiferous  areas.  Spiracles  large,  elongate- 
oval,  placed  obliquely  on  the  sides  of  a  deep  split  and  so  capable  of  close  approximation. 
Anal  gills  four,  each  short,  tapering  gradually  to  the  blunt  tip. 

Head  capsule  compact,  massive,  as  in  tribe.  Labrum  transversely  oval,  margin  with  short 
yellowish  hairs  and  a  larger  tuft  on  either  side.  Antenna  two-segmented,  second  segment 
rather  stout,  cylindrical,  slightly  arcuate;  apical  papilla  small  but  high.  Mandible  broad  and 
flattened,  with  a  small  dorsal  tooth  and  a  row  of  five  ventral  teeth.  Maxilla  generalized  in 
structure,  as  in  tribe.  Hypopharynx  as  in  Limnobaria,  consisting  of  a  roughly  circular 
chitinized  collar  provided  with  a  crown  of  stout  teeth.  Mentum  broad,  undivided,  anterior 
margin  with  eleven  teeth. 

Pupa. —  Length  (including  breathing  horns),  8-9  mm. 
Length  of  breathing  horns,  1.2-1.3  mm. 
Width  of  body,  d.-s.,  0.85-0.9  mm. 
Depth,  d.-s.,  1-1.05  mm. 

Pronotal  breathing  horns  grayish  subhyaline;  head  and  thorax  with  sheaths  dark  brown; 
abdomen  whitish,  hooks  and  spines  brown. 

Cephalic  crest  small,  indistinctly  bilobed,  not  setiferous;  front  long  and  parallel;  rostral 
sheath  very  long  and  narrow,  subtended  on  either  side  by  sheaths  of  paraglossae,  the  latter 
projecting  beyond  tip  of  rostrum  and  ending  almost  opposite  end  of  wing  sheath;  margin 
of  cheeks  flattened  as  in  Limnobaria.  Antennal  sheaths  short,  ending  slightly  beyond  base 
of  wing  pad.  Pronotal  breathing  horns  very  large  and  prominent,  not  contiguous  basally; 
about  a  dozen  breathing  pores  along  dorsal  margin.  Meson otum  unarmed;  wing  sheaths 
ending  opposite  base  of  third  abdominal  segment;  leg  sheaths  ending  opposite  or  slightly 
beyond  midlength  of  fourth  abdominal  segment;  tarsal  sheaths  ending  about  on  a  level. 
Abdominal  segments  3  to  7  near  base  with  two  bands  of  chitinized  hooks  arranged  in  curved 
transverse  rows  inclosing  an  oval  transverse  area.  Cauda  chitinized,  tergal  region  produced 
into  two  parallel  curved  hooks  bending  strongly  dorsad. 

Found  at  Alto  Pass,  Union  County,  Illinois,  June  6,  1919. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  819 

Genus  Dicranomyia  Stephens  (Gr.  fork  +fly) 

1818  Furcomyia  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  1,  p.  106  (nomen  nudum). 

1829  Dicranomyia  Steph.     Cat.  Brit.  Ins.,  vol.  2,  p.  243. 

1830  Siagona  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  6,  pi.  65,  figs.  5-7. 
1830  Glochina  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  6,  p.  280. 

1854     Numantia  Bigot.    Ann.  Soc.  Ent.  France,  ser.  3,  vol.  2,  p.  470. 

Larva.' — Form  slender.  Body  nearly  glabrous,  abdominal  and  thoracic  segments  with 
dorsal  and  ventral  transverse  welts  on  basal  annuli.  Spiracular  disk  small,  the  five  lobes 
indistinct  but  indicated,  spiracles  large.  Anal  gills  four,  slender.  Head  capsule  massive, 
of  the  Limnobia  type.  Mouth  parts  almost  as  in  Limnobia;  ventral  cutting  edge  of  man- 
dible with  fewer  teeth;  mentum  with  anterior  margin  more  transverse  and  with  fewer 
teeth. 

Pupa. —  Cephalic  crest  lacking.  Pronotal  breathing  horns  broad,  in  D.  simidans  with 
a  basal  recurved  hook  on  dorsal  side,  in  other  known  species  unarmed.  Basal  abdominal 
annuli  with  transverse  welts.  Dorsum  of  eighth  abdominal  segment  with  vestigial  spiracles. 

Dicranomyia  is  a  very  extensive  genus  including  more  than  two  hun- 
dred described  species  of  usually  small  flies  which  are  found  in  most  parts 
of  the  world.  The  immature  stages,  which  are  found  in  a  variety  of 
habitats  practically  as  extensive  as  is  covered  by  the  entire  family  of 
crane-flies,  range  from  forms  that  are  almost  strictly  aquatic,  thru  species 
living  beneath  the  bark  of  trees,  to  still  other  species  which  are  leaf  miners. 

In  Europe,  Dicranomyia  trinotata  (Meig.)  is  a  characteristic  member 
of  the  hygropetric  association,  the  insects  living  on  rocks  in  streams, 
where  they  are  covered  with  a  thin  sheet  of  water  and  are  usually  asso- 
ciated with  such  insect  forms  as  Beraea,  Tinodes,  Stactobia  (Trichoptera), 
Orphnephila  testacea  (Ruthe),  Pericoma  nubila  (Meig.),  Dixa  maculata 
Meig.,  Oxycera  pulchella  Meig.,  and  other  Diptera.  The  larva  is  cylin- 
drical, measuring  from  10  to  11  millimeters  in  length  and  from  1.5  to 
2  millimeters  in  diameter.  The  dorsal  surface  is  greenish  mottled  with 
darker,  the  ventral  surface  brighter.  The  larvae  live  in  loosely  spun 
silken  cases  in  which  they  pupate.  The  pupae  are  about  10  millimeters 
long,  and  live  in  cocoons  which  are  almost  horizontal  in  position.  The 
mature  pupa  breaks  thru  the  cocoon  by  means  of  its  sharp-edged  breathing 
horns,  the  adult  then  creeping  forth  and  leaving  the  cast  pupal  hull  behind. 
(Thienemann,  1909:64-65,  and  Grtinberg,  1910:29.) 

Dicranomyia  dumetorum  Meig.  lives  in  decaying,  principally  deciduous, 
wood.  Winnertz  (1853)  found  it  in  large  numbers  in  a  decaying  beech 
tree,  associated  with  Bremia  cilipes  (Winn.). 


820  CHARLES  PAUL  ALEXANDER 

V 

A  species  of  crane-fly  doubtfully  referred  to  Dicranomyia  pilipennis 
Egg.  (Schubart,  1854)  has  been  found  in  ditch  water  in  Holland. 

Dicranomyia  umbrata  de  Meij.,  a  Javan  species,  lives  in  the  slimy  green 
algae  floating  in  stagnant,  as  well  as  flowing,  water.  The  pupae  live  in 
cocoons  in  the  algal  sheath,  with  the  cephalic  end  projecting.  The  larvae 
are  from  10  to  12  millimeters  in  length,  cylindrical,  about  0.6  millimeter 
in  diameter,  and  of  a  yellowish  color.  The  head  is  almost  entirely 
retractile.  The  body  is  almost  smooth,  having  only  an  inconspicuous 
transverse  welt  on  the  sscond  abdominal  segment  near  the  posterior 
margin.  The  caudal  end  is  somewhat  enlarged  and  is  truncated  behind. 
De  Meij  ere  (1916:197-198)  supplies  a  good  description  of  the  structure 
of  the  larval  head  capsule. 

Dicranomyia  foliocuniculator  Swez.,  of  the  Hawaiian  Islands,  is  the 
only  recorded  leaf-mining  crane-fly.  It  was  found  by  Swezey  mining 
in  the  leaves  of  a  species  of  Cyrtandra  (Gesneriaceae)  in  the  island  of 
Oahu.  Pupation  of  the  species  takes  place  within  .the  mines  (Swezey, 
1913  and  1915). 

In  North  America  a  number  of  species  are  known.  Dicranomyia 
simulans  has  been  ably  discussed  by  Needham  (1908 a: 214-217)  and  later 
by  Malloch  (1915-17 b  [1917]).  This  species  is,  for  the  most  part,  a 
member  of  the  hygropetric  fauna,  dwelling  in  usually  lotic  water  where 
it  is  associated  with  a  characteristic  rapid-stream  fauna.  Other  con- 
ditions under  which  the  species  is  found  are  discussed  later.  D.  badia 
and  D.  siulta  live  in  and  under  saturated  moss  cushions.  D.  macateei 
Alex,  has  been  bred  from  larvae  in  decaying  wood  (Dr.  W.  G.  Dietz). 
D.  rara  O.  S.  has  been  bred  from  larvae  in  a  rotten  willow,  the  larvae 
being  taken  on  Plummers  Island,  Maryland,  by  H.  S.  Barber  on  Octo- 
ber 12,  1913,  and  emerging  as  adults  on  November  14. 

Dicranomyia  simulans  (Walk.) 

1848    Limnobia  simulans  Walk.     List  Dipt.  Brit.  Mus.,  vol.  1,  p.  45. 
The  best  account  of  the  life  history  of  the  common  and  widespread 
Dicranomyia  simulans   is    that   by   Needham    (1908 a: 214-2 16),    quoted 
below : 

It  is  abundant  on  the  piers  along  the  west  shore  of  Lake  Michigan.  .  .  .  This  pier 
[at  Lake  Forest,  Illinois]  was  built  on  heavy  driven  piling,  covered  outside  with  heavy  plank. 
About  three  feet  of  surface  was  exposed  above  the  water  at  its  normal  stage.  The  planks 
were  old,  and  sheltered  a  scanty  growth  of  short,  stemmed  mosses  in  the  cracks,  and  bore 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  821 

a  heavy  fringe  of  Cladophora  and  other  algae  just  below  the  water  line,  with  a  film  of  "  skin 
algae  "  extending  a  little  higher. 

All  over  the  sides  of  the  plank,  in  either  sun  or  shade,  the  adult  simulans  could  be  seen 
throughout  the  summer  months,  sometimes  in  considerable  numbers.  I  was  first  attracted 
to  notice  them  by  their  habit  of  running  rapidly  sidewise  along  the  pier,  and  their  resemblance 
to  harvestmen  (Phalangidae).  They  run  habitually  sidewise,  apparently  rarely  moving 
forward  except  to  escape  an  obstruction,  and  very  rarely  appearing  on  the  top  of  the  pier. 
They  rest  in  an  inverted  position  on  the  under  surface  of  the  overhanging  plank  on  the  top 
of  the  pier.  They  stiok  to  the  surface  so  persistently  that  it  is  difficult  to  make  one  take 
flight;  they  may  be  driven  all  about  on  the  surface,  or  poked  with  a  stick;  they  can  fly  well 
enough  when  they  will,  but  when  induced  to  fiy  they  settle  again  almost  at  once,  and  within 
a  few  feet  of  their  starting  place. 

They  are  associated  upon  the  piers  with  Geranomyia  canadensis  and  with  numerous 
midges  and  micro-caddis  flies  (Hydroptilidae)  and  a  few  larger  caddis  flies  of  the  genus 
Hydropsyche. 

Males  are  more  in  evidence,  but  probably  not  more  abundant  in  fact.  The  females  come 
out  from  their  resting  places  only  to  lay  their  eggs,  and  are  only  to  be  seen  when  busily  engaged 
in  the  performance  of  this  task.  They  stand  on  tiptoe,  with  the  long  ovipositor  held  in 
vertical  position  at  the  tip  of  the  deflexed  abdomen,  and  they  swing  the  body  up  and  down 
in  rapid  shuttlelike  vibration,  freely  rising  and  falling  on  the  long  and  widely  outspread 
legs.  Thus  the  point  of  the  ovipositor  is  driven  against  the  wet  surface  of  the  plank,  thrusting 
almost  as  rapidly  as  the  needlebar  of  a  sewing  machine;  it  is  moved  about  over  the  surface, 
as  if  searching  for  soft  spots  in  the  wood,  and  occasionally  it  makes  a  deeper  thrust  when  a 
suitable  place  is  found,  and  an  egg  is  deposited. 

The  egg-laying  process  is  often  interrupted  and  is  continuously  interfered  with  by  the 
too  importunate  males.  When  a  male  in  running  about  on  the  plank  comes  upon  a  female 
ovipositing,  he  stands  directly  above  her  at  the  full  upward  stretch  of  his  legs,  while  she 
goes  right  along  with  her  work;  but  the  instant  she  ceases  her  vibrating  and  lifts  her  ovi- 
positor, he  is  ready  with  his  forceps,  upturned  and  outspread  at  the  tip  of  his  decurved  abdo- 
men, to  seize  her.  Usually  she  does  not  want  to  be  interrupted  and  moves  away,  while  he 
tries  to  run  parallel  and  maintain  all  the  while  his  position  of  vantage  above  her.  Often 
other  males  are  encountered,  and  then  the  males  engage  in  a  rough  and  tumble  fight.  They 
push  and  shove  each  other  in  a  most  ludicrous  manner,  reminding  one  of  pigs  fighting, 
and  often  an  encounter  of  this  sort  enables  the  female  to  escape  and  go  on  quietly  with 
her  work. 

The  males  have  well  developed  eyes,  but  their  sight  must  be  very  poor;  for,  while  always 
searching  for  females,  they  seem  quite  unable  to  find  them  by  sight,  often  passing  females 
at  work  within  a  distance  of  a  few  centimeters.  But  their  tactile  sense  seems  more  acute, 
When  a  male  in  running  to  and  fro  had  passed  several  times  within  six  centimeters  of  a 
female  without  noticing  her,  was  deflected  from  his  course  toward  her  by  an  obstruction 
I  purposely  placed  in  his  way,  he  instantly  sprang  toward  her  upon  the  slightest  contact, 
even  of  tips  of  tarsi,  but  was  quite  unheeding  until  this  contact  occurred.  If  it  did  not  occur 
he  would  pass  on,  even  by  the  narrowest  margin. 

All  stages  are  found  together  on  the  piers.  The  eggs  are  laid  in  the  soft  spots  in  the  old 
wood,  where  the  surface  of  the  pier  is  kept  wet,  but  not  continually  covered  by  water,  in 
the  zone  of  the  "  skin  algae."  The  larvae  live  exposed  or  thinly  algae  covered,  and  crawl 
about  slowly  over  the  wet  surface.  They  are  greenish  in  color  and  very  inconspicuous. 
In  a  cavity  among  the  stems  of  the  dwarf  mosses  (Bryum  binum  Schoeb.  var.  varium  Lindb. 
and  Amblystegium  orthocladon  Lesq.  and  James)  in  a  crevice  at  the  upper  limit  of  the 
wet  area  the  larva  spins  about  itself  a  sheet  of  tissue  and  fastens  bits  of  moss  stems  and 
leaves  to  its  outside,  and  transforms  inside  the  tube  thus  formed  into  a  pupa.  The  tube 
is  longer  than  its  body,  and  the  pupa  moves  in  or  out  at  will,  doubtless  by  the  aid  of  the 
hooks  at  the  ends  of  its  body. 


822  CHARLES  PAUL  ALEXANDER 

The  following  descriptions  of  the  immature  stages  are  adapted  from 
those  of  Needham  and  Malloch: 

Larva. —  Length,  10-15  mm. 
Diameter,  1.5-2  mm. 

Coloration  green,  with  distinct  brown  marks  on  dorsum  and  with  an  interrupted  mid- 
dorsal  row  of  alternating  paler  dots  and  crossmarks;  dark  area  made  up  of  closely  placed 
spinous  hairs,  clear  areas  for  the  most  part  devoid  of  hairs. 

Body  cylindrical,  abruptly  tapering  behind  on  last  abdominal  segment.  Legs  lacking,  but 
a  scurfy,  pubescent  creeping-fold  on  under  surface  of  meso-  and  metathorax,  a  similar  one  on 
first  abdominal  segment,  and  much  larger,  transversely  placed,  fusiform  creeping-ridges  on 
ventral  surface  of  abdominal  segments  2  to  7,  on  anterior  ring  of  each  segment.  Incisions 
between  dorsal  segments  of  abdomen  margined  with  blackish  spinules,  which  are  slightly 
stronger  than  other  hairs  of  dorsum.  Spiracular  disk  vertically  cleft,  with  sloping  sides, 
folded  together  when  under  water,  border  of  aperture  fringed  with  short  hairs  and  destitute 
of  fleshy  lobes.  Spiracles  oval.  Anal  gills  four,  fleshy. 

Head  large,  similar  in  general  appearance  to  that  of  Limnobia,  entirely  retractile  within 
enlarged  prothorax;  head  showing  a  broad,  pale  yellow,  median. band,  sides  black  from  base 
of  antennae  backward.  Labrum  transversely  oval,  with  a  margin  of  close-set,  scurfy  hairs; 
clypeus  one-fourth  broader  than  labrum,  yellow,  with  parallel  sides  but  emarginate  on  front 
for  reception  of  labrum ;  three  recurved,  stout  setae  on  lateral  margin  of  clypeus  on  each  side, 
one  on  each  angle  and  two  on  disk.  Mentum  slightly  convex  in  outline,  median  tooth  much 
longer  and  stouter  than  first  lateral,  second-  and  third  laterals  as  large  as  median  tooth. 
Antenna  long,  the  shaft  about  three  times  as  long  as  its  greatest  diameter.  Maxillary  palpi 
short  and  inconspicuous. 

Pupa. —  Length,  8-9  mm. 
Diameter,  1.5  mm. 

Body  smooth  and  shiny,  ends  brownish.  Front  of  thorax  upcurved  dorsally.  Pronotal 
breathing  horns  broad,  laterally  flattened,  obtuse  at  apex,  each  with  a  basal  recurved,  sharp 
hook  on  'its  dorsal  side;  breathing  tubercles  arranged  in  a  semicircular  row  along  obtuse  tip 
of  horns.  Dorsum  of  thorax  with  a  faint  fretwork  of  raised  lines  on  surface.  Abdomen 
smooth,  with  transverse  lines  of  scurfy  pubescence,  terminating  in  a  pair  of  stout,  sharply 
recurved  hooks. 

Malloch  (1915-17  b,  pi.  33,  fig.  5)  has  figured  the  peculiar  pronotal 
breathing  horn  of  this  species. 

Dicranomyia  stulta  0.  S. 

1859    Dicranomyia  stulta  0.  S.    Proc.  Acad.  Nat.  Sci.  Phila.,  p.  210. 

Adults  of  Dicranomyia  stulta  are  often  exceedingly  abundant,  flying 
about,  or  resting  in  close  proximity  to,  rocky  ledges  or  cliffs  near  streams. 
They  are  found  commonly  in  June,  associated  with  such  crane-fly  species 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  823 

as  Geranomyia  canadensis,  Dactylolabis  montana,  Tipula  ignobilis,  T.  api- 
calis,  and  similar  forms.  The  larvae  live  in  and  beneath  the  saturated 
cushions  of  moss  (Amblystegium)  that  grow  on  the  shale  near  the  water's 
edge.  The  only  associated  crane-fly  larvae  found  near  Cascadilla  Creek, 
Ithaca,  New  York,  where  this  species  is  common,  were  Tipula  ignobilis, 
the  larvae  of  both  species  being  exceedingly  abundant. 

Larva.—  Length,  10-12.2  mm. 
Diameter,  0.6-0.7  mm. 

Color  pale  whitish  with  a  green  cast;  transverse  abdominal  welts  dark  brown. 

Form  rather  long  and  slender.  In  addition  to  dorsal  and  ventral  welts  on  abdominal 
segments  2  to  8,  a  complete  band  at  base  of  metathorax  and  ventral  bands  on  mesothorax 
and  first  abdominal  segment;  ventral  abdominal  bands  larger  and  more  conspicuous  than 
narrow  dorsal  welts.  A  few  erect  setae  on  body.  Spiracular  disk  (Plate  XXVII,  105)  rather 
small,  with  a  deep  vertical  split,  the  large  ovate  spiracles  capable  of  close  approximation; 
ventral  lobes  blunt,  with  a  black  spot  on  face,  fringed  with  short  black  hairs  and  with  two 
sensory  setae;  each  blunt  lateral  lobe  narrowly  lined  with  a  black  crescent;  dorsal  lobes  very 
small,  dusky;  spiracular  disk  fringed  with  short,  dark  hairs.  Anal  gills  four,  large  and  pale. 

Head  capsule  as  in  tribe.  Labrum  (Plate  XXVII,  106)  subtriangular;  anterior  margin 
broad,  nearly  straight  across,  with  a  dense  fringe  of  hairs  which  are  coarser  at  ends  of 
lobe;  near  anterior  margin  of  labrum  two  oval,  hyaline  areas,  each  with  three  short  papil- 
lae; a  few  sensory  setae  along  anterior  margin.  Mentum  (Plate  XXVII,  107)  broad, 
anterior  outline  triangular,  running  out  into  a  rather  long  apical  point,  each  side  with  about 
five  teeth.  Hypopharynx  (Plate  XXVII,  108)  as  in  this  group  of  genera,  consisting  of  a 
collar  of  two  chitinized  combs,  each  with  about  ten  sharp  teeth.  Antenna  (Plate  XXVII,  109) 
short,  cylindrical,  the  apical  papilla  very  small,  reduced  to  a  tiny  disk.  Mandible  (Plate 

XXVII,  110)  broad,  flattened,  with  a  blunt  apical  point  which  is  only  a  little  longer  than 
the  teeth  on  either  side  of  it;  ventral  cutting  edge  with  about  five  blunt  teeth,  which  are 
gradually  smaller  from  the  outermost  toward  the  base;  inner  face  of  mandible  with  a  blunt 
prosthecal  tooth  and  an  oblique  fringe  of  coarse  setae.     Maxilla  (Plate  XXVII,  111)  with 
the  cardines  large,  with  about  three  setiferous  punctures;  stipites  short,  cylindrical;  outer  lobe 
fringed  with  long  hairs  and  bearing  the  short,  flattened,  disklike  palpus,  which  has  five  or  six 
hyaline  pegs;  inner  lobe  smaller,  with  dense,  short  hairs  and  a  few  sensory  organs. 

Pupa. —  Length,  about  6  mm. 

Labrum  very  obtusely  rounded  at  apex,  not  bilobed.  Labial  lobes  straight  across  or  very 
slightly  convex  across  posterior  margin.  Maxillary  palpi  narrowed  toward  tips  (Plate 

XXVIII,  112).     Pronotal  breathing  horns   (Plate  XXVIII,   113  and   114)   elongate-oval, 
earlike,  the  ventral  margin  more  bulging,  the  apex  a  little  narrowed  but  obtuse;  a  row  of 
breathing  pores  along  outer  margin,  beginning  on  lateral  face  near  dorsal  margin,  these  few 
in  number  and  widely  separated,  becoming  more  numerous  toward  apex  of  organ.     Leg 
sheaths  as  usual  in  this  group  of  genera,  those  of  fore  legs  the  longest,  those  of  hind  legs 
the  shortest.     Male  cauda  (Plate  XXVIII,  115)  with  ventral  lobes  (Plate  XXVIII,  116) 
large,  bluntly  rounded  at  tips;  two  small,  brown,  approximated  tubercles  at  base  of  split  on 


824  CHARLES  PAUL  ALEXANDER 

ventral  side;  two  blunt  tubercles  on  dorsal  surface  near  base  and  close  to  median  line  (Plate 
XXVIII,  117).  Tergal  lobes  at  outer  angles  of  a  flattened  plate,  very  short,  triangular,  each 
with  two  small  hairs  on  caudal  face  before  tip. 

Nepiorwtype  —  Cascadilla  Creek,  Ithaca,  New  York,  May  22,  1917. 

Neanotype. —  Type  locality,  June  6,  1917. 

Paratypes. —  Abundant  larvae  and  pupae  with  types,  May  22  to  June  6,  1917. 

Dicranomyia  badia  (Walk.) 

1848    Limnobia  badia  Walk.     List  Dipt.  Brit,  Mus.,  vol.  1,  p.  46. 

1859     Dicranomyia  humidicola  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  210. 

Dicranomyia  badia  is  a  very  common  species  thruout  eastern  North 
America,  occurring  in  gorges  and  rayines  and  along  streams.  The  adult 
flies  may  be  found  resting  on  perpendicular  cliffs  near  these  haunts.  In 
a  position  of  rest  they  have  all  six  feet  on  the  support,  a  very  different 
resting  position  from  that  of  the  often-associated  genera  Oropeza  and 
Dolichopeza. 

The  larvae,  as  is  frequent  in  this  genus,  live  in  and  beneath  moss, 
especially  saturated  cushions  of  moss  growing  in  or  near  the  margins  of 
streams.  They  are  of  a  clear  light  pea-green  color,  and  simulate  the  moss 
to  an  astonishing  degree.  The  writer  found  these  larvae  in  Needham's 
Glen,  Ithaca,  New  York,  on  April  16,  1917,  beneath  wet  cushions  of 
Amblystegium  irriguum  (Hook.  &  Wils.)  B.  &  S.,  a  moss  that  covers 
all  the  rocks  and  stones  near  water.  In  these  moss  cushions  the  larvae 
were  associated  with  tiny  larvae  of  Tipula  ignobilis  and  pupae  of  T. 
collaris.  When  ready  to  pupate,  the  larva  spins  a  small,  silken,  silt- 
covered  case,  which  is  further  protected  by  a  covering  of  small  pieces  of 
moss  stems  adhering  to  its  outside,  this  case  being  hung  up  in  the  moss 
cushion,  with  the  cephalic  end  of  the  pupa  projecting.  The  pupal  duration 
is  about  seven  days  (April  21  to  28  in  the  cases  observed).  The  dark- 
colored  pupa,  with  its  contrasting  yellow  breathing  horns,  is  very 
handsome. 

Larva. —  Length,  10.5  mm. 

Diameter,  0.6-0.65  mm. 

Coloration  light  pea-green,  abdominal  welts  brown;  after  death  general  color  fading  to 
very  pale  greenish  white. 

Form  slender;  body  terete,  dorsal  and  ventral  transverse  welts  at  base  of  abdominal  seg- 
ments conspicuous.  Spiracular  disk  (Plate  XXVII,  104)  with  lobes  indistinct,  suffused  with 
dusky,  lateral  lobes  very  blunt;  disk  surrounded  by  a  fringe  of  short,  dark-colored  hairs. 
Spiracles  very  large,  ovate,  dorsal  ends  close  together.  Anal  gills  long  and  slender,  pale. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  825 

Head  capsule  and  mouth  parts  similar  to  those  of  D.  stulta,  already  described,  judging  from 
the  scanty  material  of  D.  badia  available  for  study. 

Pupa. —  Length,  8-8.5  mm. 

Width,  d.-s.,  1.1  mm. 
Depth,  d.-v.,  1.2  mm. 

Head,  thorax,  and  appendages  dark  brown;  pronotal  breathing  horns  light  yellow;  abdomen 
greenish,  the  cauda  chitinized,  light  brown. 

Labrum  very  broad,  indistinctly  bilobed  at  tip.  Labial  lobes  large,  broadly  transverse, 
posterior  margin  almost  straight  across.  Maxillary  palpi  broad,  tips  truncated  (Plate XXIX, 
119).  Lateral  margins  of  cheeks  flattened  into  ledges. 

Pronotal  breathing  horns  large,  flattened,  in  lateral  outline  (Plate  XXIX,  118)  subcircular  or 
nearly  so,  with  a  row  of  rather  widely  separated  breathing  tubercles  along  margin;  as  viewed 
from  above,  horns  directed  proximad,  so  as  to  be  contiguous  at  tips.  A  high  median  crest 
on  mesonotum  behind  breathing  horns.  Wing  sheaths  ending  before  apex  of  abdominal 
segment  2.  Leg  sheaths  ending  far  before  apex  of  abdominal  segment  4;  as  usual  in  this 
division,  the  hind  legs  a  little  the  shortest,  the  fore  legs  a  little  the  longest.  Abdominal 
segments  with  a  distinct  basal  welt  which  is  thickly  margined  with  microscopic  curved  hooks. 
Lateral  spiracles  distinct,  but  small  and  probably  nonfunctional.  Female  cauda  with  sternal 
valves  shorter  than  long  tergal  valves,  the  latter  (Plate  XXIX,  120)  almost  straight,  each  with 
a  powerful,  acute  spine  on  lateral  margin  at  about  midlength,  this  directed  dorsad.  Near  the 
margin  of  segment  8,  on  dorsum,  a  pair  of  rudimentary  spiracles. 

Nepionotype  —  Needham's  Glen,  Ithaca,  New  York,  April  16,  1917.  (No.  5-1917.) 
Neanotype. —  Type  locality,  May  7,  1917. 

Genus  Rhipidia  Meigen  (Gr.  a  fan) 

1818    Rhipidia  Meig.    Syst.  Beschr.  Zweifl.  Ins.,  vol.  1,  p.  153. 
1911     Ceratostephanus  Brun.     Rec.  Indian  Mus.,  vol.  6,  p.  271. 

Larva.' —  Form  rather  stout,  body  terete.  Abdominal  sternitea  1  to  7  and  tergites  2  to 
7  with  narrow  transverse  basal  welts  of  chitinized  points.  Spiracular  disk  with  indistinct 
lobes.  Head  capsule  massive,  not  unlike  that;  of  Dicranomyia.  Labrum  broadly  transverse. 
Mandible  very  broad,  flattened,  with  only  three  ventral  cutting  teeth.  Maxilla  of  simple 
structure.  Antenna  with  apical  papilla  or  segment  very  flattened,  disklike.  Hypopharynx 
of  two  chitinized  plates,  each  with  about  twelve  comblike  teeth.  Mentum  almost  transverse 
across  anterior  margin,  with  from  nine  to  eleven  teeth,  the  outermost  fused. 

Pupa.—  Pronotal  breathing  horns  elongate  for  this  subtribe,  about  three  times  as  long  as 
broad.  Abdomen  with  transverse  bands  of  spicules  on  tergites  3  to  7  and  sternites  5  to  7, 
and  on  extreme  lateral  parts  of  sternites  3  and  4. 

Rhipidia  is  a  small  to  medium-sized  genus  (about  thirty-five  species) 
having  its  center  of  distribution  in  the  American  tropics,  with  some 
species  occurring  thruout  temperate  Europe  and  America  and  a  less  number 
in  Africa  and  the  Oriental  region.  The  genus  is  based  on  a  sexual  char- 


826  CHAELES  PAUL  ALEXANDER 

acter,  the  pectinate  antennae  of  the  male,  and  several  of  the  species 
run  inconveniently  close  to  Dicranomyia. 

The  immature  stages  are  spent  beneath  the  bark  of  decaying  trees  or 
in  decaying  vegetable  or  animal  matter.  In  Europe  the  genotype, 
Rhipidia  maculata  Meig.,  has  been  recorded  as  living  in  old  cow-manure. 
Beling  found  the  insects  in  such  a  situation,  associated  with  the  larvae 
of  Rhyphus  pundatus  (Fabr.)  (Beling,  1879:52-53)  and  a  staphylinid 
beetle,  Platystethus  morsitans  Payk.  (Beling,  1873  b:  592).  R.  uniseriata 
Schin.  was  found  by  Beling  (1879:53-54)  living  in  decaying  beech  wood, 
in  company  with  larvae  of  Xylota  lenta  Meig.  and  X.  segnis  (Linn.)  (Syr- 
phidae)  as  well  as  with  larvae  of  a  tipuline  crane-fly,  Ctenophora.  The 
larvae  of  this  species,  like  those  of  R.  maculata,  live  in  thin  silken  cases, 
open  at  both  ends  and  covered  with  particles  of  wood  and  other  debris. 

In  America,  besides  Rhipidia  bryanti  (which  is  discussed  in  some  detail) 
the  following  records  of  the  immature  stages  are  available:  Rhipidia 
maculata,  recorded  in  Europe  as  living  in  decaying  organic  matter,  was 
found  by  Needham  (1908 a:  170,  204)  in  tent  traps  set  over  the  bed  of 
Beaver  Meadow  Brook  in  the  Adirondacks,  the  insects  presumably  hav- 
ing emerged  from  the  stream  bed  or  from  the  thin  layers  of  moss  covering 
the  exposed  stones.  R.fidelis  O.  S.  was  reared  from  larvae  in  decaying  wood 
near  Ithaca,  New  York,  by  Carl  Ilg.  This  species  belongs  to  the  same 
subgenus  (Monorhipidia  Alex.)  as  the  European  R.  uniseriata,  which  has 
similar  larval  habits.  R.  domestica  0.  S.  has  been  bred  from  larvae 
obtained  in  fermented  sap  of  the  sour  gum  (Nyssa  sylvatica  Marsh., 
Cornaceae)  at  Clementon,  New  Jersey  (Johnson,  1910:704).  Males  and 
females  of  this  species  were  bred  by  Popenoe  at  Washington,  D.  C., 
from  larvae  on  more  or  less  decaying  roots  of  taro  (Colocasia  antiquorum 
Schott,  Araceae)  taken  at  Gough,  South  Carolina,  on  February  1,  1911. 
Specimens  of  R.  domestica  emerged  on  August  21,  1906,  at  Juneau, 
Alaska,  in  radishes  infested  with  H-ylemyia  brassicae  (Bouche) . 

Rhipidia  (Rhipidia)  bryanti  Johns. 

1909    Rhipidia  bryanti  Johns.    Proc.  Boston  Soc.  Nat.  Hist.,  vol.  34,  p.  123-124,  pi.  16, 
fig.  20. 

Rhipidia  bryanti  is  one  of  the  largest  and  handsomest  species  of  the 
genus,  and  is  widely  distributed  thruout  the  eastern  United  States  from 
Maine  to  Texas.  The  immature  stages  are  spent  beneath  the  decaying 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  827 

bark  of  trees.  The  material  studied  was  collected  by  J.  R.  Malloch  in 
Potomac  Park,  Washington,  D.  C.,  on  May  11,  1913,  and  was  reared 
by  R.  C.  Shannon  (No.  35-1913),  whose  notes  on  the  subject  are  as  follows: 

A  small  colony  (about  seven)  of  tipulids  were  found  in  a  hollow  of  a  tree  behind  the  bark. 
The  larvae  were  covered  with  slime,  which  gathered  debris  as  they  moved  about  and  so 
formed  a  case.  The  next  morning  one  had  pupated.  Two  larvae  were  boiled  and  pre- 
served in  alcohol,  while  two  others  were  preserved  in  formal.  As  they  move  about  they 
leave  a  path  of  slime  behind  them.  On  May  14,  the  one  that  had  pupated  on  the  12th 
emerged.  On  May  18  another  imago  issued. 

There  is  an  obvious  error  in  the  duration  of  the  pupal  stage  as  given 
• —  only  two  days.  It  is  possible  that  a  pupa  was  included  in  the  material 
and  was  overlooked  in  its  case  of  debris.  The  date  of  emergence  of  the 
second  adult  would  show  a  pupal  duration  of  six  days,  which  is  probably 
nearly  correct. 

Larva. —  Length,  13.2-14  mm. 
Diameter,  1  mm. 

Coloration  white. 

Form  terete,  rather  stout.  Sternites  1  to  7  and  tergites  2  to  7  with  narrow  transverse  welts 
on  basal  rings;  welts  yellow,  provided  with  long,  transverse  rows  of  microscopic  roughened 
points,  those  along  margins  of  welts  coarser,  those  in  centers  very  tiny;  these  rows  inter- 
rupted along  pleura.  Caudal  end  blunt,  with  lobes  of  spiracular  disk  indistinct.  Spiracles 
large,  capable  of  close  approximation. 

Head  capsule  massive,  of  the  Limnobiatype.  Labrum  (PlateXXX,  121)  broadly  transverse, 
anterior  margin  almost  straight  across,  near  margin  two  oval  areas  which  are  provided  with 
small  sensory  papillae.  Epipharyngeal  region  with  abundant  hairs.  Mentum  (Plate  XXX, 
122)  chitinized,  the  anterior  margin  almost  transverse,  with  a  large  median  tooth  and  about 
three  lateral  teeth  on  either  side,  the  outermost  of  these  an  evident  fusion  of  about  three  lesser 
teeth.  Hypopharynx  (PlateXXX,  123  and  124)  asusualin  this  subtribe,  a  collar-like  structure 
composed  of  two  parallel  combs  united  at  the  ends;  the  larger  plate  broadly  elongate,  surface 
with  abundant  scalelike  plates,  anterior  margin  with  twelve  teeth,  the  lateral  ones  narrow, 
the  teeth  gradually  enlarging  toward  the  middle,  the  middle  pair  a  little  shorter;  second  plate 
of  hypopharynx  broadly  transverse,  narrow,  likewise  with  about  twelve  teeth,  the  two  outer 
most  on  each  side  long,  subacute,  the  middle  tooth  shorter  and  more  flattened;  between  the 
two  middle  teeth  a  small  triangular  or  conical  point.  Antenna  (Plate  XXX,  125)  short  cylin- 
drical, the  apical  papilla  a  very  flattened  disk  or  button  which  is  much  broader  than  long. 
Mandible  (Plate  XXX,  126)  very  broad,  flattened,  with  the  apical  tooth  rather  long  and  slen- 
der; two  smaller  teeth  dorsad  and  three  others  ventrad,  the  dorsal  teeth  blunt,  the  ventral 
teeth  truncated,  the  most  basal  tooth  very  broad.  Maxilla  (Plate  XXX,  127)  of  the  simple 
generalized  structure  of  this  tribe;  cardines  large,  with  two  setiferous  punctures;  stipites 
weakly  chitinized  basally;  outer  lobe  of  maxilla  with  the  large  flattened  palpus  at  its  tips; 
palpus  surrounded  by  numerous  long  setae  and  having  several  hyaline  sense  pegs  at  its 
apex;  inner  lobe  of  maxilla  smaller,  with  numerous  long  hairs,  especially  a  slitlike  brush 
near  margin. 


828  CHARLES  PAUL  ALEXANDER 

Pupa.- —  Length,  about  12  mm.  (cast  skin). 

Labrum  broad,  rounded  at  apex.  Labial  lobes  broadly  transverse,  caudal  margin 
indistinctly  trilobed.  Maxillary  palpi  large,  flattened,  apex  bluntly  pointed;  a  rather  angular 
tooth  on  margin  near  base  (Plate  XXXI,  128).  Cheeks  produced  into  flattened  ledges. 
Antennae  short  and  stout. 

Pronotal  breathing  horns  (Plate  XXXI,  129)  elongate  for  this  subtribe,  about  three  times 
as  long  as  the  greatest  diameter,  flattened,  apical  half  slightly  expanded,  margin  with  tiny 
tubercles.  Wing  sheaths  attaining  base  of  abdominal  segment  3.  Leg  sheaths  attaining 
base  of  abdominal  segment  5;  tarsi  ending  about  on  a  level,  or  hind  tarsi  a  little  the  shorter 
and  fore  tarsi  a  little  the  longer.  Abdomen  with  basal  bands  of  setae  on  tergites  3  to  7 
and  sternites  5  to  7,  and  on  the  extreme  lateral  parts  of  sternites  3  and  4;  these  bands  thickly 
margined  with  tiny,  golden-yellow  hairs  or  points,  the  median  part  of  each  band  naked  or 
nearly  so.  Male  cauda  (Plate  XXXI,  130)  with  dorsal  lobes  (Plate  XXXI,  131)  very  small, 
more  or  less  flattened,  divergent,  rather  blunt  at  tips;  ventral  lobes  elongate,  contiguous 
along  inner  face. 

Nepionotype.—  Potomac  Park,  D.  C.,  May  11,  1913. 
Neanotype. —  Cast  pupal  skin,  wi€h  type  larva,  May  14,  1913. 
Paratype. —  One  larva  with  type. 

Subtribe  Dicranoptycharia 

The  subtribe  Dicranoptycharia,  so  far  as  known,  includes  only  the 
genus  Dicranoptycha.  The  division  is  close  to  the  Rhamphidaria  but 
is  easily  separated  from  it  in  all  stages. 

Genus  Dicranoptycha  Osten  Sacken  (Gr.  fork  +  fold) 

1818    Marginomyia  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  1,  p.  147  (nomen  nudum). 
1859     Dicranoptycha  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  217. 

Larva. —  Form  very  elongate,  terete.  Integument  smooth,  glassy,  transparent. 
Abdominal  segments  2  to  8  each  with  a  basal  transverse  band  or  area  of  microscopic  chitinized 
points  on  ventral  surface;  segment  8  with  a  similar  band  on  dorsum.  Spiracular  disk  sur- 
rounded by  four  lobes,  the  lateral  pair  more  slender  than  the  blunt  ventral  pair;  dorsal  lobe 
very  low  or  lacking;  a  triangular  brown  mark  on  disk  between  spiracles.  Spiracles  small, 
widely  separated.  Anal  gills  a  fleshy  protuberant  ring  surrounding  anus. 

Head  capsule  compact,  massive,  the  prefrons  large  with  a  few  marginal  punctures;  externo- 
lateral  plates  very  broad.  Labrum  large,  flattened,  pale.  Antenna  two-segmented;  apical 
segment  almost  as  long  as  basal  segment,  gradually  narrowed  to  the  blunt  tip.  Mandible 
with  a  blunt  dorsal  and  two  blunt  ventral  teeth.  Maxilla  generalized  in  structure.  Hypo- 
pharynx  a  rounded  cushion.  Mentum  deeply  split  behind  but  not  completely  divided, 
with  three  principal  teeth  and  a  small  reduced  lateral  tooth  on  either  side. 

Pupa.' —  Cephalic  crest  low,  depressed,  setiferous.  Labrum  tumid.  Labial  lobes  oval, 
contiguous.  Antennal  sheaths  ending  opposite  base  of  wing  pad.  Pronotal  breathing 
horns  microscopic,  represented  only  by  tiny  triangular  tubercles.  Mesonotum  unarmed. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  829 

Wing  sheaths  ending  opposite  middle  of  third  abdominal  segment.  Leg  sheaths  ending 
opposite  base  of  fifth  abdominal  segment,  the  tarsi  terminating  on  a  level  or  nearly  so. 
Abdominal  tergites  and  sternites  each  with  four  transverse  rows  of  microscopic  setae;  lateral 
spiracles  on  segments  2  to  7;  no  apparent  spiracles  on  dorsum  of  segment  S. 

Dicranoptycha  is  a  principally  Holarctic  genus  including  twelve  known 
species,  six  of  which  occur  in  the  United  States.  The  flies  are  of  medium 
size  and  of  a  dull,  inconspicuous  appearance,  and  are  rather  difficult  of 
taxonomic  separation.  The  life  histories  of  members  of  this  genus  have 
only  recently  been  ascertained.  The  following  notes  are  taken  from 
more  detailed  accounts  in  earlier  papers  by  the  writer  (Alexander,  1919, 
a  and  b). 

The  habits  of  the  immature  stages  of  Dicranoptycha  may  be  briefly 
summarized  as  follows:  The  larvae  and  the  pupae  live  in  rich  humous 
soil  overlain  with  a  cover  of  leaf  mold  and  other  vegetable  debris.  They 
frequent  open  woods  where  there  is  more  or  less  shubbery  and  tall  herbage. 
Running  streams  or  rills  are  not  necessary  for  the  development  of  the 
immature  stages.  The  larvae  live  in  the  uppermost  zone  of  the  soil,  where 
they  are  associated  with  a  rather  characteristic  group  of  animal  forms, 
such  as  dipterous  larvae  (Sciara),  beetle  larvae,  and  centipedes.  They 
are  characterized  by  the  exceedingly  long,  slender  body  and  the  shiny 
glabrous  skin,  and  may  be  confused  with  no  other  dipterous  larva  yet 
made  known.  The  glassy  appearance  of  the  body  suggests  the  shiny 
shell  of  a  small  univalve  snail,  the  dead  fragments  of  which  occurred  in 
some  numbers  in  the  same  situations.  The  larvae  of  Dicranoptycha 
winnemana  Alex.,  an  Austral  species,  attain  a  length  of  from  20  to  22 
millimeters,  with  an  average  diameter  of  only  1  millimeter.  The  larvae 
of  the  genus  are  herbivores.  When  ready  to  pupate  they  incase  them- 
selves in  compact  earthen  cells,  which  are  10  x  3.5  millimeters  in  size, 
firm  in  texture,  and  rather  thick-walled  but  apparently  without  silk. 
There  is  a  small  opening  at  either  end.  The  pupal  period  is  about  ten 
days  or  possibly  a  little  less.  Pupation  takes  place  in  the  relatively  dry 
soil  that  forms  the  larval  haunt. 

The  pupa  of  Dicranoptycha  winnemana  measures  from  9.1  to  12.8 
millimeters  in  total  length.  The  width  at  the  wing  pad  is  about  1.7 
millimeters.  The  diagnostic  features  are  given  above  at  some  length 
for  the  genus  and  need  not  be  repeated  here.  For  other  notes  the  reader 
is  referred  to  the  papers  already  cited. 


830  CHARLES  PAUL  ALEXANDER 

The  adult  flies  of  the  various  species  of  Dicranoptycha  are  usually 
abundant  where  they  occur.  They  may  be  found  resting  on  the  upper 
surface  of  leaves  of  shrubbery  and  tall  herbage  in  open  woods,  often  far 
from  water,  which  is  not  so  necessary  for  development  as  with  most 
species  of  Tipulidae. 

Subtribe  Rhamphidaria 

The  division  Rhamphidaria  includes  the  genus  Rhamphidia,  with  the 
possible  addition  of  a  few  exotic  genera.  Rhamphidia  is  one  of  the  con- 
stituent genera  of  the  former  group  Antochini,  but  differs  considerably 
from  all  others  whose  life  histories  have  been  made  known.  The  closest 
relatives  of  Rhamphidia  seem  to  be  the  Dicranoptycharia,  on  the  one  hand, 
and  the  lowermost  divisions  of  the  Hexatomini,  on  the  other.  The 
larvae  and  the  pupae  of  Rhamphidia  present  a  curious  eriopterine  appear- 
ance, but  their  structure  indicates  only  a  distant  relationship  with  the 
Eriopterini. 

Genus  Rhamphidia  Meigen  (Gr.  rostrum) 

1825  Megarhina  St.  Farg.  et  Serv.     Encyclop.  Method.,  Ins.,  vol.  10,  part  2,  p.  585. 

1825  Helius  St.  Farg.  et  Serv.     Encyclop.  Method.,  Index,  p.  831. 

1829  Leptorhina  Steph.     Cat.  Brit.  Ins.,  vol.  2,  p.  243. 

1830  Rhamphidia  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  6,  p.  281. 

Larva. —  Body  terete.  A  transverse  welt,  covered  with  microscopic  scabrous  points,  on 
ventral  surface  of  basal  annuli  of  abdominal  segments  2  to  7.  Spiracular  disk  surrounded 
by  five  short  lobes  which  are  fringed  with  abundant,  rather  long,  hairs.  Head  capsule 
massive,  generalized  in  structure.  Mandible  short  and  stout,  ending  in  two  subequal  blunt 
teeth.  Maxilla  consisting  of  two  rather  short,  densely  hairy  lobes.  Antenna  short. 
Hypopharynx  chitinized,  outer  margin  with  about  a  dozen  teeth.  Mentum  not  completely 
divided,  with  five  teeth.  Coloration  of  body  dark  brown,  produced  by  the  dense  appressed 
pubescence  covering  it. 

Pupa.- —  Cephalic  crest  double,  the  anterior  part  low,  the  posterior  part  the  larger;  lobes 
setiferous,  divergent.  Two  setae  on  front  between  eyes.  Pronotal  breathing  horns  long 
and  slender,  slightly  curved.  Mesonotum  convex.  Wing  sheaths  reaching  end  of  second 
abdominal  segment.  Leg  sheaths  ending  just  before  posterior  margin  of  fourth  abdominal 
segment;  all  the  tarsi  about  equal  in  length,  or  those  of  fore  legs  a  little  longer.  Abdominal 
segments  with  two  narrow  basal  rings  and  a  broader  posterior  ring  having  four  narrow 
transverse  bands  of  spicules  and  comparatively  few  setae.  Spiracles  rather  large  on 
pleurites  2  to  7,  and  a  large  conspicuous  pair  on  dorsum  of  segment  8. 

Rhamphidia  is  a  small  genus  of  crane-flies  (about  thirty-five  species), 
widely  distributed  thruout  the  temperate  and  tropical  regions  of  the  world. 
The  larvae  of  the  European  R.  longirostris  Meig.  were  found  by  Gercke 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  831 

(1884)  in  submerged  stems  of  the  water  dock,  Rumex  aquaticus  Linn. 
The  eggs  are  described  as  being  rather  long,  white,  and  granulate.  It  is 
suggested  that  the  developmental  stages  may  be  associated  with  water. 
The  two  local  species  have  been  reared  and  are  discussed  herewith. 

Rhamphidia  mainensis  Alex. 

1916    Rhamphidia  mainensis  Alex.    Proc.  Acad.  Nat.  Sci.  Phila.,  p.  498-499,  fig.  14. 

Rhamphidia  mainensis  appears  to  be  a  rather  uncommon  f6rm,  much 
rarer  than  R.  flavipes,  the  other  local  species.  Adult  flies  were  not 
uncommon  in  the  Basin  Swamp,  Orono,  Maine,  on  June  12,  1913.  This 
swamp  is  a  low,  sunken  area  surrounded  on  most  sides  by  hills,  opening 
into  the  "  Basin/'  an  affluent  of  the  Penobscot  River.  Cold  springs  of 
water  percolate  down  from  these  hillsides,  and  the  soil  is  very  wet,  boggy, 
and  richly  filled  with  organic  matter.  The  chief  floral  constituents  are 
a  few  elms  and  white  birches  and  an  abundance  of  alders,  Spiraea  latifolia 
Borkh.,  and  S.  tomentosa  Linn.  The  herbage  consists  of  ferns  such  as 
Onoclea  and  Osmunda,  patches  of  Iris,  Impatiens  biflora  Walt.,  and  many 
rushes  and  sedges.  Crane-flies  associated  with  R.  mainensis  on  the 
date  mentioned  included  the  following:  Dicranomyia  haeretica, 
Epiphragma  fascipennis,  Pseudolimnophila  luteipennis,  P.  inornata, 
Limnophila  fasciolata,  L.  macrocera,  Pilaria  recondita,  Ulomorpha  pilosetta, 
Tricyphona  inconstans,  Erioptera  vespertina,  and  Tipula  sulphurea,  also 
an  abundance  of  Ptychoptera  rufocincta  and  Bittacomorpha  clavipes. 

Larvae  were  first  found  on  April  20,  1917,  in  the  dark,  cold  swamp 
known  as  Larch  Meadows,  south  of  Ithaca,  New  York.  Here  they 
occurred  in  the  thick,  black,  saturated  organic  matter  comprising  the 
soil  of  the  swamp.  The  vegetation  consists  of  the  dominant  alder  (Alnus 
incana  [Linn.]  Moench.),  the  poison  sumac  (Rhus  Vernix  Linn.),  and 
the  marsh  marigold  (Caltha  palustris  Linn.),  as  well  as  an  abundance 
of  other  plant  species  in  lesser  numbers.  The  earthy  material  in  which 
the  crane-fly  larvae  were  found  was  full  of  the  organic  remains  of  plants, 
such  as  ferns,  leaves,  alder  catkins,  and  the  like.  Associated  with  these 
larvae  at  this  time  were  numerous  small  hydrophilid  beetles,  and  a  great 
abundance  of  larvae  of  a  dascillid  beetle  of  the  subfamily  Helodinae, 
of  various  sizes  and  ages.  Numerous  tabanid  and  stratiomyiid  larvae, 
and  the  larvae  of  the  crane-fly  species  Bittacomorpha  clavipes,  Pseudolimno- 
phila luteipennis,  and  Tipula  dejecta,  also  occurred. 


832  CHAELES  PAUL  ALEXANDER 

The  larvae  were  conspicuous  by  their  dark  coloration  and,  when  placed 
in  water,  by  their  active,  snakelike  movements,  in _ this  regard  being  very 
different  from  somewhat  similar  larvae  of  certain  Eriopterini  which  they 
resemble  superficially  because  of  the  five  subequal  lobes  surrounding  the 
spiracular  disk.  The  indoor  pupal  period  is  six  days  (May  8  to  14,  1917). 

Larva. —  Length,  9-11 .2  mm. 

Diameter,  0 . 75-0 . 9  mm. 

Coloration  dark  brown,  sutures  pale;  pale  spots  on  dorsum  and  on  sides  of  body;  spiracular 
disk  pale,  lobes  marked  with  darker. 

Form  rather  stout,  terete  (Plate  XXXI,  132) .  Body  densely  covered  with  a  long,  appressed, 
dark  pubescence.  Prothoracic  segment  narrowed  in  front,  long,  divided  into  two  rings  by 
a  faint  constriction;  mesothoracic  and  metathoracic  segments  gradually  longer.  First 
abdominal  segment  shorter  than  last  thoracic  segment;  abdominal  segments  2  to  7  long, 
each  divided  into  two  narrow  annuli  by  a  deep  constriction  which  is  destitute  of  hairs  but 
has  just  before  it  a  sharp,  transverse  ridge  of  stiff  hairs;  short,  incomplete  ridges  of  these 
stiff  hairs  on  dorsum  of  anterior  ring;  anterior  ring  about  half  as  long  as  posterior  ring;  on 
ventral  side  of  each  of  segments  2  to  7  on  anterior  ring,  a  transverse  swelling,  these  swellings 
becoming  more  convex  and  prominent  on  posterior  segments;  swellings  appearing  almost 
smooth,  being  covered  only  with  microscopic,  roughened  points;  posterior  ring  with  a  sharp 
ridge  of  hairs  at  about  two-thirds  its  length.  Chaetotaxy  as  follows:  dorsal  segments  with 
short  setae  at  about  midlength  of  posterior  ring;  a  single  rather  stout,  black  bristle  on  pleura 
of  anterior  ring,  immediately  above  transverse  swellings;  a  similar  seta  on  posterior  ring; 
two  groups  of  very  long,  delicate  setae  on  ventral  face  of  posterior  ring,  one  on  each  side  of 
the  ridge  of  erect  hairs;  thoracic  segments  approximately  similar  to  abdominal  segments,  but 
sternal  setae  at  about  midlength  even  more  prominent.  Last  segment  of  body  elongated,  the 
spiracular  disk  (Plate  XXXI,  136)  surrounded  by  five  lobes;  dorsal  lobe  the  smallest,  rather 
blunt,  inner  face  with  a  brownish,  triangular-oval  mark;  lateral  lobes  of  medium  length, 
inner  face  suffused  with  brown,  which  is  darkest,  almost  black,  on  lower  edge  of  lobe;  ventral 
lobes  the  longest,  inner  face  of  each  with  two  broad,  parallel,  blackish  lines,  separated  by 
a  somewhat  narrower  pale  line;  entire  disk  fringed  around  with  long,  dark-colored  hairs, 
which  are  longest  near  tips  of  lobes,  where  they  are  strongly  recurved,  almost  pencil-like; 
fringe  continuous  between  dorsal  and  lateral  lobes,  but  between  ventral  and  lateral  lobes, 
and  between  the  two  ventral  lobes,  hairs  toward  base  of  each  lobe  very  short  to  lacking; 
ventral  lobes  just  before  tips  with  a  single  long  sensory  bristle.  Spiracles  large,  subcircular, 
situated  at  base  of  lateral  lobes,  dark-colored,  narrowly  margined  with  pale.  Anal  gills 
consisting  of  two  pairs  of  pale,  stout,  cylindrical  lobes,  tapering  toward  tips,  before  which 
there  is  a  slight  constriction. 

Head  capsule  massive  and  compact,  of  the  normal  generalized  limnobiine  type.  Labrum 
large,  conspicuous,  transverse,  densely  fringed  with  long  hairs;  on  epipharyngeal  region, 
a  large,  dense  tuft  of  moderately  elongate  hairs  on  either  side  of  median  line.  Mentum 
(Plate  XXXI,  133)  not  completely  divided,  but  with  a  very  deep  split  behind,  a  large  median 
tooth,  and  two  smaller  teeth  on^ either  side;  behind  (dorsad  of)  elongate  median  tooth,  a 
slightly  wider  flattened  lobe  whose  margins  extend  beyond  those  of  outer  tooth.  Hypo- 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  833 

pharynx  a  broad  semicircular  band  of  chitin  whose  anterior  margin  is  provided  with 
about  a  dozen  teeth,  the  intermediate  ones  more  blunt  and  rounded,  the  lateral  teeth  longer 
and  more  slender.  (The  antennae  of  this  species  were  not  distinguishable  in  the  specimens 
available,  but  are  undoubtedly  similar  to  those  of  R.  flavipes  described  hereinafter.)  Mandible 
(Plate  XXXI,  135)  of  the  generalized  limnobiine  type,  short  and  stout,  terminating  in  two 
blunt  teeth;  ventral  cutting  edge  with  about  three  teeth  which  are  gradually  smaller  toward 
base  of  mandible;  a  dorsal  row  of  two  teeth,  of  which  the  basal  one  is  the  smaller;  base  of 
mandible  on  outer  face  (heel)  prolonged  into  an  acute  flattened  blade;  a  conspicuous  tuft 
of  hairs  at  prosthecal  region.  Maxilla  consisting  of  two  rather  short,  stout  lobes  which 
are  shorter  than  the  mandible,  densely  provided  with  short  hairs;  palpi  large,  shaped  like 
a  half  of  a  short  cylinder  split  lengthwise. 

Pupa.- —  Very  similar  to  that  of  R.  flavipes,  as  described  on  the  following  pages.  (The 
writer  has  only  the  cast  pupal  skin  of  R.  mainensis,  and  it  seems  to  agree  very  closely  with 
the  pupa  of  R.  flavipes  except  that  the  lobes  of  the  labial  sheaths  are  longer  and  more  pointed, 
and  the  labrum  is  a  little  longer  to  provide  for  the  longer  rostrum  of  the  former  species.) 

Nepionotype. —  Ithaca,  New  York,  April  20,  1917. 
Neanotype. —  Cast  pupal  skin,  May  14,  1917. 

Rhamphidia  flavipes  Macq. 

1855  Rhamphidia  flavipes  Macq.     Dipt.  Exot.,  5th  supp.,  p.  17. 

1856  Rhamphidia  prominens  Walk.     Ins.  Saunders,  vol.  1,  Dipt.,  p.  435. 
1859    Rhamphidia  brevirostris  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  222. 

Rhamphidia  flavipes  is  a  common  and  widely  distributed  fly  thruout 
the  eastern  United  States  and  Canada.  The  species  is  characteristic 
of  cat-tail  swamps  and  similar  situations.  It  has  been  reared  from  leaves 
of  bur  reed,  Sparganium,  brought  in  by  C.  H.  Kennedy  from  Ringwood 
Hollow,  near  Etna,  New  York,  in  September.  These  larvae  were  asso- 
ciated with  larvae  of  Prionocera  fuscipennis,  likewise  a  characteristic 
inhabitant  of  open  swamps.  The  specimens  here  described  were  taken 
in  a  small  cat-tail  swamp  near  Bool's  hillside,  Ithaca,  New  York,  in  June, 
1917.  Here  they  were  associated  with  a  number  of  larvae  of  characteristic 
swamp-inhabiting  crane-flies,  such  as  Ptychoptera  rufocincta,  Pseudolimno- 
phila  luteipennis,  Limnophila  macrocera,  Pilaria  recondita,  Liogma  nodi- 
cornis  (in  moss) ,  Prionocera  fuscipennis,  and  Tipula  tricolor. 

This  is  unquestionably  the  larva  that  was  found  by  Hart  and  doubtfully 
referred  by  him  to  the  genus  Erioptera  (Hart,  1898  [1895] :  197-199,  also 
Malloch,  1915-17  b :  237).  Later,  Mik  (1898 : 62)  doubted  that  this  belonged 
to  Erioptera  because  of  the  long  lashes  of  hairs  surrounding  the  caudal 
lobes,  a  character  not  shown  by  typical  eriopterine  larvae.  The  larvae 
that  Hart  found  were  living  among  rushes  and  other  vegetation  floating 


834  CHARLES  PAUL  ALEXANDER 

on  the  surface  of  the  water  in  the  Illinois  River.     When  submerged  these 
larvae  were  very  active,  quite  as  noted  in  the  account  of  R.  mainensis. 

Larva.' —  In  all  general  features  like  larva  of  R.  mainensis  (with  the  material  available 
the  writer  is  unable  to  point  out  differences).  Present  species  a  little  larger  than  R.  mainensis, 
measuring  from  12  to  13  mm.  in  length  and  1  mm.  in  diameter.  Mouth  parts  and  head 
capsule  almost  exactly  like  those  of  R.  mainensis,  already  described.  Antenna  (Plate  XXXI, 
134)  with  first  segment  elongate,  a  little  enlarged  toward  tip,  and  with  an  elongate,  thimble- 
shaped  papilla  at  tip  whose  surface  is  delicately  sculptured;  surrounding  this  papilla  at  tip 
of  basal  segment,  a  few  microscopic  tubercles  and  pegs;  on  face  of  basal  segment  on 
proximal  half,  a  circular  porous  plate. 

Pupa.—  Length,  7.8-8.8  mm. 
Width,  d.-s.,  1  mm. 
Depth,  d.-v.,  1.1-1.2  mm. 

Coloration  light  brown;  abdomen  paler,  trivittate  with  dark  brown;  a  broad,  dorso-median 
line,  and  narrower,  somewhat  interrupted,  pleural  stripes;  breathing  horns  yellow,  darker 
at  extreme  bases. 

Cephalic  crest  (Plate  XXXII,  137)  low,  the  lobes  divergent,  their  lateral  angles  with  a  stout 
seta  which  is  directed  cephalad.  Just  behind  the  anterior  crest  a  much  larger,  low,  appressed 
lobe,  which  is  transversely  wrinkled,  bearing  on  its  side  a  seta  directed  dorsad.  Eyes  rather 
large,  widely  separated  by  front;  two  setae  on  front  between  eyes.  Labrum  triangular, 
pointed  at  apex.  Front  a  little  elongated  to  provide  for  short  rostrum  of  adult.  Sheaths 
of  maxillary  palpi  long,  stout,  almost  straight.  Labial  palpi  contiguous,  lying  side  by  side 
at  tip  of  labrum.  Antennae  rather  widely  separated  at  their  bases,  ending  just  beyond 
wing  root;  in  males  lying  across  face  of  eye,  due  to  the  large  size  of  eyes  in  this  sex. 

Mesonotum  prominent,  carinate  medially  (Plate  XXXIII,  142).  Breathing  horns  long  and 
slender,  slightly  curved,  transversely  crenulated,  a  little  enlarged  toward  tips;  proximo- 
cephalad  of  base  of  each  horn  a  small  rounded  tubercle  bearing  two  setae.  Lateral  angles 
of  thorax  with  about  three  short  setae.  Mesonotum  convex,  transversely  wrinkled,  with 
a  few  very  short  setae.  Wing  sheaths  ending  about  opposite  posterior  margin  of  second 
abdominal  segment.  Leg  sheaths  parallel,  about  subequal  in  length  or  those  of  fore  legs  a 
little  longer,  ending  just  before  posterior  margin  of  fourth  abdominal  segment.  Dorsal 
abdominal  segments  (Plate  XXXIII,  143)  each  with  two  narrow  basal  rings  and  a  much 
broader  posterior  ring;  ventral  segments  (Plate  XXXIII,  144)  with  the  two  basal  rings 
confluent;  dorsal  segments  with  each  narrow  basal  ring  having  a  narrow,  slightly  arcuated 
band  of  spicules,  and  posterior  ring  having  a  narrow  basal  band  of  spicules  which  is  more 
or  less  arcuated,  and  a  straight  caudal  band,  immediately  in  front  of  which  are  four  groups 
of  two  setae  each,  or,  in  some  specimens,  with  the  intermediate  groups  reduced  to  a  single 
seta;  ventral  segments  with  posterior  ring  having  a  broad  transverse  band  of  spicules, 
with  three  setae  in  alinement  at  each  end  of  this  band;  closer  to  base  of  segment,  two 
strong  setae  on  either  side;  anterior  ring  darkened.  Pleura  with  large  but  apparently 
nonfunctional  spiracles  on  segments  2  to  7,  a  strong  ante-spiracular  seta,  and  three  post- 
spiracular  setae,  as  illustrated  (Plate  XXXIII,  144).  Male  cauda  (Plate  XXXII,  139  and 
1^0)  terminating  in  two  acute  tips  which  are  directed  dorsad;  at  base  of  these  lobes  on 


THE  CKANE-FLIES  OF  NEW  YORK  —  PART  II  835 

dorsal  face,  two  blunt,  approximated,  parallel  lobes  directed  cephalad;  at  the  base  of  long 
apical  lobes,  a  shorter,  pointed,  setiferous  lobe,  directed  laterad  and  dorsad,  bearing  on  its 
side  and  near  the  apex  a  few  setae.  Dorsum  of  segment  8  with  four  prominent  lobes 
surrounding  a  hollow,  these  lobes  ending  in  acute  tips,  the  posterior  pair  directed  more  dorsad, 
the  anterior  pair  directed  more  laterad,  at  the  base  of  the  latter  a  pair  of  spiracles.  Female 
cauda  similar  but  with  acidothecae  prolonged,  bearing  on  the  side  prominent  lateral  lobes 
(Plate  XXXII,  141),  corresponding  to  the  setiferous  lobe  of  male  as  described  above; 
quadrangle  of  tubercles  on  dorsum  of  segment  8  (Plate  XXXII,  138)  about  as  in  male,  all 
the  tubercles  ending  in  chitinized  points. 

Nepionotype. —  Ithaca,  New  York,  May  11,  1917. 
Neanotype. —  Ithaca,  New  York,  June  3,  1917. 
Paratypes. —  Pupae  with  neanotype. 

Tribe  Hexatomini 

The  tribe  Hexatomini,  as  here  understood,  comprises  a  very  extensive 
group  of  medium-sized  crane-flies,  made  up  of  the  members  of  the  former 
groups  Limnophilini  and  Hexatomini.  The  more  generalized  members 
of  the  group  are  herbivorous,  but  the  two  highest  subtribes,  Limnophilaria 
and  Hexatomaria,  are  carnivorous  in  their  feeding  habits. 

The  larvae  of  the  lower  divisions  have  the  head  capsule  massive  and 
compact,  the  mentum  chitinized  and  with  the  anterior  margin  toothed, 
and  the  hypopharynx  usually  well  preserved.  The  mandibles  are  not 
formed  into  long,  curved  hooks,  and  the  maxillae  are  of  a  generalized 
organization.  The  higher  subtribes,  Limnophilaria  and  Hexatomaria, 
have  the  head  capsule  long  and  slender  and  very  much  dissected,  with 
the  constituent  plates  very  narrow  and  separate.  The  mentum  is  very 
reduced  or  lacking,  in  some  of  the  Limnophilaria  consisting  of  an  articulated 
transverse  bar  which  is  grooved  with  parallel  fine  striae.  In  the  group 
Ulomorphae  and  in  the  Hexatomaria  the  chitinized  mental  region  is 
nearly,  if  not  quite,  lacking,  allowing  for  great  distention  of  this  region 
of  the  head.  The  mandibles  are  powerful,  curved  hooks,  bearing  a  few 
teeth  at  about  midlength.  In  the  group  Ulomorphae  the  mandibles 
are  hinged  at  about  midlength,  the  basal  part  being  deeply  grooved  on 
the  inner  face  to  receive  the  blade  in  a  position  of  rest.  The  maxillae 
have  the  outer  lobe  prolonged  into  an  elongate  flattened  blade  which 
extends  out  of  the  thoracic  orifice  when  the  head  is  retracted. 

The  pupae  of  the  Ularia  and  the  Epiphragmaria  have  a  pair  of  large 
spiracles  on  the  dorsum  of  the  eighth  abdominal  segment,  indicating  a 
close  phylogenetic  relationship  with  the  Limnobiini.  Dactylolabis  has 


836  CHARLES  PAUL  ALEXANDER 

the  lateral  abdominal  spiracles  protuberant,  with  those  of  the  second 
segment  very  large  and  conspicuous.  Pseudolimnophila,  the  Ulomorphae, 
and  Pilaria  have  the  pronotal  b'reathing  horns  very  long  and  slender, 
with  the  tips  split  into  divergent  flaps.  In  the  Hexatomaria  the  lateral 
abdominal  spiracles  are  large  and  functional. 

The  immature  stages  of  the  Polymeraria,  including  the  single  tropical 
American  genus  Polymera  Wiedemann,  are  unknown.  Their  discovery 
might  result  in  the  inclusion  of  this  sub  tribe  with  the  Pediciini  rather 
than  with  the  Hexatomini. 

The  following  keys  separate  the  subtribes  of  the  tribe  Hexatomini: 

Larvae 

1.  Men  turn  completely  divided,  each  half  with  seven  teeth  on  anterior  margin;  hypopharynx 

a  semicircular  chitinized  ring  with  numerous  teeth  around  anterior  margin. 

Pseudolimnophilaria  (p.  848) 
Mentum  when  present  not  divided;  hypopharynx  not  shaped  as  above 2 

2.  Abdominal  segments  2  to  7  with  a  basal  transverse  creeping-welt 3 

Abdominal  segments  2  to  7  without  such  a  welt 4 

3.  Mentum  three-toothed;  antenna  elongate,  the  apical  segment  hemispherical. 

Epiphragmaria  (p.  843) 
Mentum  nine-toothed;  antenna  short,  subglobular,  the  apex  with  two  short  papillae. 

Ularia  (p.  838) 

4.  Body  depressed;   head  capsule  massive,   compact;  mentum  heavily  chitinized,   seven- 

toothed;  maxilla  not  projecting Dactylolabaria  (p.  852) 

Body  terete;  head  capsule  of  slender  bars,  not  compact;  mentum  feeble,  at  most  a  narrow, 
transverse  bar  which  is  delicately  striate;  maxilla  projecting  from  thoracic  orifice 5 

5.  Mental  bar  present Limnophilaria  (group  -Limnophilae)  (p.  858) 

Mental  bar  lacking 6 

6.  Dorsal  plates  of  head  capsule  firmly  united;  epipharynx  and  maxilla  densely    hairy; 

mandible  hinged Limnophilaria  (group  Ulomorphae)  (p.  869) 

Dorsal  plates  of  head  capsule  widely  separated,   at  most  merely  contiguous    behind; 
epipharynx  and  maxilla  not  hairy;  mandible  not  hinged Hexatomaria  (p.  876) 

Pupae 

1.  Two  large  spiracles  on  dorsum  of  eighth  abdominal  segment 2 

No  large  spiracles  as  above ! 3 

2.  Pronotal  breathing  horns  subchitinized,  directed  strongly  ventrad;  abdominal  segments 

without  shagreened  transverse  bands;  pupates  in  wood Epiphragmaria  (p.  843) 

Pronotal  breathing  horns  flattened,  directed  laterad;  shagreened  transverse  bands  on 
abdominal  segments;  pupates  in  earth Ularia  (p.  838) 

3.  Abdominal  segments  with  large  protuberant  spiracles,  those  on  second  segment  very 

large Dactylolabaria  (p.  852) 

Abdominal  segments  without  conspicuous  protuberant  spiracles 4 

4.  Pronotal  breathing  horns  elongate,  split  into  two  flaps  at  tip 5 

Pronotal  breathing  horns  not  split  into  two  such  flaps 6 

5.  Abdominal  segments  with  five  or  six  rows  of  setiferous  tubercles. 

Pseudolimnophilaria  (p.  848) 
Abdominal  segments  without  such  rows  of  tubercles. 

Limnophilaria  (group  Ulomorphae}  (p. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  837 

6.  Head  and  thorax  without  spines  or  tubercles;  lateral  abdominal  spiracles  small. 

Limnophilaria  (group  Limnophilae)  (p.  858) 

Head  and  thorax  often  with  tubercles  on  antennal  scape,  labrum,  or  mesonotal  scutellum; 
lateral  abdominal  spiracles  large,  functional Hexatomaria  (p.  876) 

The  most  important  literature  on  the  tribe  Hexatomini  is  as  follows: 

Ula  macroptera Larva Stannius,  1829: 205. 

Ula  macroptera Larva,  pupa,  general.  .  Ferris,  1849:337-341. 

Ula  macroptera- General Alexander,  1915 a: 2. 

Ula  macroptera General Pierre,  1919-20:76. 

Ula  bolitophila General Loew,  1869: 4-5. 

Ula  elegans Larva,  pupa,  general. .  .  Alexander,  1915  a. 

Ula  elegans Larva,  pupa Malloch,  1915-17 b: 226.  (Copy.) 

Epiphragma  picta Larva Bremi-Wolf ,  1846. 

Epiphragma  picta Larva,  pupa.. Beling,  1873 b : 589-590. 

Epiphragma  fascipennis Larva,  pupa,  general...    Needham,  1903:281-285. 

Epiphragma  fascipennis Larva,  pupa Malloch,  19 15-17 b: 224-225. 

(Copy.) 

Pseudolimnophila  luteipennis Larva,  pupa,  general. . .  Hart,  1898  [1895]:  202-204. 

Pseudolimnophila  luteipennis Larva,  pupa,  general. .  Malloch,  19 15-17 b: 222-223. 

Dactylolabis  wodzickii Larva,  pupa,  general.  .   Nowicki,  1867:340-343. 

Dactylolabis  denticulata Larva,  pupa,  general .  .   Mik,  1894. 

Limnophila  ferruginea Larva,  pupa,  general.  .   De  Meijere,  1916:204-206. 

Limnophila  hyalipennis Larva,  pupa Beling,  1886: 198-199. 

Limnophila  lineola Pupa , .  .  .  Beling,  1879:54. 

Limnophila  lineola *.  .  .  .   Larva Beling,  1886: 199-200. 

Limnophila  nemoralis Larva,  pupa Beling,  1886:200-201. 

Limnophila  ochracea General Beling,  1886: 202. 

Limnophila  pallida Larva,  pupa Beling,  1873  a:  556-558. 

Limnophila  dispar Larva,  general Perris,  1849:331. 

Limnophila  pictipennis Larva,  pupa,  general..  .   Beling,  1879:51-52. 

Limnophila  pictipennis Larva Brauer,  1883: 55. 

Limnophila  punctata Larva Scheffer,  in  Rossi,  1848: 10. 

Limnophila  punctata Larva Osten  Sacken,  1869:201. 

Limnophila  punctata Larva,  pupa,  general.  .   Beling,  1886: 195-197. 

Limnophila  punctata Larva Gerbig,  1913: 158-161. 

Limnophila  punctata General Cameron,  1917:63. 

Limnophila  bryobid. General Mik,  1881 : 205-206. 

Limnophila  sinistra Larva,  pupa Hudson,  1920: 33-34. 

Pilaria  fuscipennis Larva,  pupa Beling,  1886: 197-198. 

Pilaria  fuscipennis Larva Gerbig,  1913: 164-166. 

Pilaria  fuscipennis General Cameron,  1917: 63. 

Pilaria  discicollis Larva Gerbig,  1913: 163-164. 

Pilaria  discicollis General Cameron,  1917: 63. 

Pilaria  tenuipes Pupa Hart,  1898  [1895] :  204-205. 

Pilaria  tenuipes Pupa Malloch,  19 1 5-17 b: 223-224. 

Eriocera spinosa Larva,  pupa,  general.  .  Alexander  and  Lloyd,   1914: 27- 

30. 

Eriocera  spinosa General Alexander,  1915  c:  149. 

Eriocera  cinerea Larva Alexander  and  Lloyd,  1914:21- 

23  (as  longicornis). 

Eriocera  longicornis *. . ,        .  Pupa,  general Alexander  and  Lloyd,   1914 : 23- 

27. 


838  CHARLES  PAUL  ALEXANDER 

Eriocera  longicornis General Alexander,  1915  c:  149-152. 

Eriocera  fultonensis Larva,  pupa,  general.  .  Alexander  and  Lloyd,  1914:30- 

33. 

Hexatoma  nigra General Von  Roser,  1834.  . 

Hexatoma  megacera Larva,  pupa,  general.  .   Alexander,  1915 c:  141-148. 

Penthoptera  albitarsis Larva,  pupa,  general.  .  Alexander,  1915 c:  152-157. 

Subtribe  Ularia 

The  division  Ularia  includes  only  the  genus  Ula.  It  represents  a  very 
primitive  group  of  crane-flies,  presumably  the  most  generalized  of  the 
entire  tribe.  The  head  capsule  of  the  larva  is  oval  and  very  massive, 
with  the  prefrons  large  and  distinct.  The  head  capsule  and  its  arrange- 
ment of  setae  is  not  conspicuously  unlike  that  of  the  eucephalous  families 
of  crane-flies,  and  this  group  of  Tipulidae  is  presumably  not  very  different 
from  the  early  tipulid  ancestors.  The  mentum  is  heavily  chitinized,  not 
completely  divided  behind,  consisting  of  two  plates,  one  behind  the  other. 
The  outermost  plate  terminates  in  three  teeth,  while  the  second  plate 
furnishes  three  additional  teeth  on  each  side.  The  hypopharynx  is  not 
chitinized.  The  antennae  are  very  small;  the  basal  segment  is  nearly 
globular,  bearing  at  its  tip  two  blunt,  conical  papillae.  The  mandibles 
are  slender;  the  ventral  cutting  edge  has  about  five  narrow  teeth;  there 
is  a  distinct  brush  of  hairs  at  the  prosthecal  region.  The  maxillae  are 
of  a  generalized  type,  with  the  palpus  large,  flattened,  and  disklike.  The 
abdomen  is  provided  with  six  creeping- welts  on  the  ventral  surface  of  the 
segments.  The  spiracular  disk  is  moderate  in  size,  squarely  truncated, 
surrounded  by  five  subequal  lobes  which  are  heavily  marked  with  black 
on  their  inner  faces. 

The  pupa  has  the  cephalic  crest  small  and  provided  only  with  very 
small  setae.  The  pronotal  breathing  horns  are  very  long,  tapering  to 
the  subacute  tips.  The  abdominal  segments  have  a  basal  transverse  band 
of  a  shagreened  appearance.  The  dorsal  spiracles  on  the  eighth  abdominal 
segment  are  large  and  distinct. 

The  closest  relative  of  the  division  is  apparently  the  genus  Epiphragma 
in  the  subtribe  Epiphragmaria.  But  this  entire  group  of  genera  (those 
included  in  the  subtribes  Ularia,  Epiphragmaria,  and  Pseudolimnophilaria) 
is  not  far  removed  from  the  tribe  Limnobiini. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  839 

Genus  Ula  Haliday  (Gr.  so/0 

1833     Ula  Hal.     Ent.  Mag.,  vol.  1,  p.  153. 

1864    Macroptera  Lioy.     Atti  dell'  Institut  Veneto,  ser.  3,  vol.  9,  p.  224. 

Larva.^-  Body  covered  with  a  short  pubescence.  Basal  annuli  of  abdominal  segments 
2  to  7  with  a  transverse  creeping-welt  on  ventral  surface.  Spiracular  disk  squarely  trun- 
cated, surrounded  by  five  subequal  lobes  which  are  fringed  with  very  short  hairs.  Head 
capsule  massive,  prefrontal  sclerite  very  large,  tapering  to  a  point  behind.  Labrum  large, 
conspicuous,  with  tufts  of  hairs.  Mandible  slender;  ventral  cutting  edge  with  five  teeth; 
a  tuft  of  hairs  on  prosthecal  region.  Maxilla  simple;  palpus  large.  Antenna  very  small; 
basal  segment  subglobular,  with  two  apical  sensory  papillae.  Mentum  not  deeply  divided 
behind;  outer  plate  with  three  apical  teeth;  inner  plate  adding  three  additional  teeth  to  each 
side.  Hypopharynx  not  chitinized. 

Pupa.- — Cephalic  crest  small,  setae  tiny.  Pronotal  breathing  horns  long,  tapering  to 
subacute,  flattened  apices.  Wing  sheaths  ending  opposite  base  of  third  abdominal  seg- 
ment. Leg  sheaths  ending  at  about  midlength  of  sixth  abdominal  segment.  Abdominal 
segments  on  tergites  and  on  apical  sternites  with  transverse  bands  of  microscopic  points, 
producing  a  shagreened  appearance;  dorsal  spiracles  on  segment  8  conspicuous. 

The  genus  Ula  includes  only  six  described  species,  of  which  three  are 
European,  two  are  North  American,  and  one  is  Javanese. 

The  adult  flies  of  the  American  species  are  commonest  in  spring  and 
late  summer.  They  frequent  cool,  shaded  gorges  and  ravines,  or  dark 
woods  in  mountainous  regions,  and  may  be  swept  from  beds  of  low  vege- 
tation, such  as  ferns,  yew,  and  other  species. 

The  larvae  of  all  the  known  species  live  in  various  species  of  fungi, 
but  go  to  earth  for  pupation. 

In  Europe  the  common  genotype,  Ula  macroptera  (Macq.),  has  been 
discussed  rather  frequently  in  the  literature.  Stannius  (1829:205)  found 
the  larvae  in  a  species  of  Agaricus.  Ferris  (1849:337-341)  furnishes  a 
brief  account  of  the  species,  from  which  the  following  notes  are  taken: 
The  fungus  in  which  the  larvae  were  found  was  Hydnum  erinaceum  Bull, 
growing  on  the  trunks  of  living  oak  trees.  The  larvae  are  gregarious 
and  frequent  galleries  in  the  fungus,  along  which  they  progress  by  means 
of  their  mandibles,  by  their  ambulatory  feet,  and  by  the  short  hairs  that 
cover  the  body.  They  were  found  in  the  month  of  November.  A  month 
later  they  went  into  the  earth,  where  they  transformed  as  pupae.  The 
pupae  bear  a  strong  resemblance  to  those  of  Limnophila,  but  differ  in 
the  shape  of  the  pronotal  breathing  horns.  In  February  and  March  the 
pupae  come  to  the  surface  of  the  earth  and  the  adults  emerge.  The 


840  CHARLES  PAUL  ALEXANDER 

caudal  end  of  the  larva  is  described  as  having  but  four  lobes;  no  mention 
is  made  of  the  median  dorsal  lobe  found  in  the.  American  species,  and  it 
is  presumably  lacking  or  very  reduced  in  size.  Pierre  (1919-20:76)  has 
reared  this  species  from  larvae  living  in  Russula  nigricans  Fries. 

Another  European  species,  U.  bolitophila  Loew,  was  bred  from  larvae 
living  in  fungi  on  beech  trees  in  Austria  (Loew,  1869:4-5). 

In  America,  the  immature  stages  of  Ula  elegans  have  been  discussed 
by  the  writer  (Alexander,  1915 a),  the  notes  given  below  being  in  part 
supplementary  to  his  earlier  account. 

Ula  elegans  0.  S. 

1869      Ula  elegans  0.  S.    Mon.  Dipt.  N.  Amer.,  part  4,  p.  276-277. 

Ula  elegans  is  a  rather  common  species  in  mountainous  regions  thruout 
the  northeastern  United  States.  The  adult  flies  swarm  in  early  spring 
and  again  in  the  fall.  At  Ithaca,  New  York,  on  May  14,  1912,  the  writer 
found  them  swarming  at  half  past  four  in  the  afternoon.  There  were 
about  fifteen  to  twenty  individuals  in  a  swarm  within  a  foot  or  two  of 
the  ground.  In  some  cases  the  swarms  were  reduced  to  two  or  three 
individuals,  or  in  a  few  instances  to  a  single  specimen.  In  copulation 
the  flies  rest  on  the  upper  surface  of  near-by  plants  (Symplocarpus,  in  the 
instance  cited),  with  all  the  legs  on  the  support.  Copulation  is  rather 
firm,  and  the  insects  fly  for  short  distances  still  united.  The  only  crane-fly 
associated  with  Ula  at  that  time  was  Limnophila  ultima  0.  S. 

On  September  15,  1912,  the  writer  found  a  fleshy  species  of  fungus 
(Fomes)  growing  on  a  much-decayed  stump  close  to  the  ground.  This 
fungus  contained  a  number  of  crane-fly  larvae,  including  about  thirty-five 
larvae  of  Limnobia  cinctipes,  a  lesser  number  of  L.  triocellata,  and  many 
larvae  of  Ula  elegans.  The  larvae  of  these  species  frequented  the  upper 
layers  of  the  mushroom  and  had  reduced  the  surface  to  a  semi-liquid 
state.  At  the  end  of  a  week  the  whole  fungus  was  reduced  to  a  very 
decayed  condition.  The  fungus,  which  was  taken  at  Gloversville,  New 
York,  was  then  transferred  to  Ithaca,  and  was  placed  in  large  glass  jars, 
with  sand  in  the  bottom  to  take  up  the  liquids  produced  by  decay  and 
to  provide  a  place  for  pupation.  The  first  adults  emerged  on  October 
14.  Emergence  continued  until  the  27th,  when  the  remaining  pupae 
were  killed  and  placed  in  alcohol.  At  that  time  they  were  very  dark- 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  841 

colored  and  evidently  nearly  ready  to  emerge  to  the  adult  state.  At 
times  the  larval  movements  are  very  active  and  eel-like,  but  at  other 
times  they  are  very  slow  and  sluggish.  At  each  movement  forward, 
the  terminal  segment  partly  telescopes  into  the  subterminal  and  is  thrown 
violently  backward.  After  transforming  to  the  adult  condition,  the  pupal 
skin  is  left  adhering  to  the  sand,  with  the  posterior  half,  or  a  smaller 
portion,  attached,  often  standing  quite  perpendicular  to  the  surface  and 
very  conspicuous. 

Larva. —  Length,  8.5-11.9  mm. 
Diameter,  1.4-1.8  mm. 

Color  dull  white;  head  capsule  very  dark  brownish  black. 

Form  moderately  slender  (Plate  XXXIV,  145),  but  the  body  not  greatly  elongated  as  in 
the  Eriopterini  and  some  Limnobiini.  Integument  covered  with  a  fine,  short  pubescence. 
Abdominal  segments  2  to  7  each  divided  by  a  constriction  into  a  narrow  basal  annulus  and 
a  broad  posterior  annulus;  basal  ring  at  about  midlength  and  at  the  end  with  narrow  trans- 
verse rows  of  tiny  setae;  abdominal  segments  2  to  7  on  ventral  surface  of  basal  ring  with 
conspicuous  raised  transverse  creeping-welts  which  are  covered  with  microscopic  points; 
eighth  abdominal  segment  suddenly  constricted  before  spiracular  disk.  Region  around  anus 
protuberant,  this  evidently  being  an  aid  to  propulsion.  Spiracular  disk  (Plate  XXXIV, 
153)  surrounded  by  five  blunt  lobes,  of  which  the  dorsal  one  is  median  in  position  and 
blunter  than  the  others;  inner  face  of  each  lobe  with  a  conspicuous  brownish  black  mark; 
a  fringe  of  delicate  blackish  hairs  surrounding  disk,  these  hairs  a  little  longer  and  more 
prominent  at  ends  of  lobes.  Spiracles  almost  circular,  widely  separated,  the  distance  between 
them  being  about  three  times  diameter  of  one  spiracle.  Disk  between  spiracles  with  an 
indistinct  arcuated  line. 

Head  capsule  (Plate  XXXIV,  146)  massive  and  compact,  rather  narrow,  posterior  incisions 
not  extending  deeply  into  capsule.  Prefrons  very  large,  tapering  gradually  to  a  sharp  point 
behind;  numerous  setae  on  prefrons  and  on  other  sclerites  of  capsule.  Labrum  (Plate  XXXIV, 
147)  large,  conspicuous;  epipharynx  and  lateral  margins  strengthened  by  narrow  bands 
of  chitin;  lateral  and  anterior  margins  of  labrum  with  a  dense  fringe  of  long  hairs;  an  irregular 
tubercle  on  dorsal  surface  on  either  side  of  labrum;  epipharynx  with  a  narrow  transverse 
band  of  setae;  juncture  of  clypeus  and  labrum  with  four  setiferous  punctures,  two  on  either 
side,  the  posterior  pair  a  little  the  closer  together;  clypeal  region  with  a  dark  transverse  basal 
and  terminal  band;  base  of  clypeus  with  three  setiferous  punctures  on  either  side.  Mentum 
(Plate  XXXIV,  148)  almost  completely  undivided,  the  usual  median  split  obliterated  except 
behind;  outermost  mental  plate  terminating  in  three  apical  teeth,  the  median  one  of  which  is 
a  little  shorter  and  broader  than  the  other  two;  behind  (dorsad  of)  outer  plate,  another  similar 
plate  which  adds  three  more  teeth  to  each  side  of  mentum.  Hypopharynx  not  chitinized. 
Antenna  (Plate  XXXIV,  149)  very  small,  short,  cylindrical  or  subglobular,  about  as  broad 
as  long;  at  the  tip  two  hyaline  papillae  shaped  like  immature  mushrooms,  the  outer  one 
about  twice  the  size  of  the  inner  one.  Mandible  (Plate  XXXIV,  150  and  151)  slender,  ending 
in  a  blunt,  slightly  curved,  apical  point;  on  ventral  cutting  edge  a  row  of  about  five  flattened 


842  CHARLES  PAUL  ALEXANDER 

teeth;  the  more  apical  ones  larger,  the  basal  one  very  broad  and  truncated;  two  or  three 
dorsal  teeth,  gradually  lessening  in  size  from  tip  of  mandible  toward  base;  two  conspicuous 
tufts  of  long  setae  at  prosthecal  region;  base  of  mandible  on  dorsal  side  with  a  large,  some- 
what curved,  plate  overlying  head  sclerites.  Maxilla  (Plate  XXXIV,  152)  with  cardo  long 
and  narrow,  transverse,  with  three  conspicuous  setiferous  punctures  bearing  long  setae; 
outer  lobe  short  and  stout,  a  little  narrowed  to  tip;  palpus  rather  large,  flattened,  disklike, 
the  truncated  apex  with  several  tiny  hyaline  pegs;  inner  lobe  with  a  sensory  bristle  and 
an  inner  fringe  of  rather  short  hairs. 

Pupa. —  Length,  6.4-7  mm. 

Width,  d.-s.,  1.2-1.3  mm. 
Depth,  d.-v.,  1.2-1.6  mm. 

Mouth  parts,  wing  sheaths,  and  leg  sheaths  rather  dark  brown;  thoracic  dorsum  and 
abdomen  light  yellowish  brown;  pronotal  breathing  horns  dark  brown,  passing  into  light 
yellow  on  apical  third  or  quarter;  mesonotal  prescutum  retaining  its  light  color  even  in  old 
pupae  and  those  preserved  in  alcohol. 

Cephalic  crest  (Plate  XXXV,  155)  represented  only  by  a  small  bilobed  protuberance 
behind  and  between  antennal  bases,  each  lobe  tipped  with  a  very  tiny  seta.  Labrum  short, 
bluntly  rounded  at  tip.  Labial  lobes  appearing  as  a  large,  roughly  quadrate  plate,  a  little 
narrowed  behind  and  with  all  angles  rounded.  Sheaths  of  maxillary  palpi  very  long  and 
slender,  rather  stout  at  base,  tapering  to  blunt  tip.  Antennae  not  conspicuous,  rather 
widely  separated  at  bases,  ending  just  before  or  opposite  origin  of  wing  pad. 

Pronotal  breathing  horns  (Plate  XXXV,  154  and  155)  long  and  conspicuous,  cylindrical, 
gradually  tapering  to  subacute  and  flattened  apices;  horns  directed  laterad  and  cephalad, 
widely  divergent.  Thoracic  dorsum  without  lobes  or  spines.  Wing  sheaths  extending  to 
just  beyond  end  of  second  abdominal  segment.  Leg  sheaths  extending  to  about  midlength 
of  sixth  abdominal  segment;  tarsal  sheaths  ending  almost  on  a  common  level,  the  fore  legs 
being  a  very  little  the  longest,  the  middle  legs  a  little  shorter.  Abdominal  tergites  with  trans- 
verse rows  of  scattered  punctures;  on  tergites  2  to  6  a  transverse  band  of  subchitinized  points, 
producing  a  shagreened  appearance;  these  bands  located  on  basal  rings  of  segments  excepting 
the  last  two,  which  are  on  extreme  ends  of  segments  5  and  6;  on  sternites  the  bands  appearing 
only  on  segments  7  and  8,  the  other  sternites  being  largely  concealed  by  the  unusually  long 
leg  sheaths;  band  on  segments  very  broad,  but  narrowly  interrupted  medially.  Male  cauda 
with  ventral  lobes  bluntly  rounded  and  inclosing  pleural  appendages  of  adult  (in  fully 
colored  pupae  these  show  thru  the  pupal  skin  as  from  twelve  to  fifteen  strong,  chitinized 
points  on  the  inner  posterior  face) ;  dorsal  lobes  (Plate  XXXV,  157)  very  short  and  blunt,  and 
closely  approximated  medially;  on  eighth  tergite  a  broad  rectangle  of  five  lobes,  including 
an  anterior  and  a  posterior  lobe  on  either  side  and  a  much  broader  anterior  median  lobe; 
just  proximad  of  anterior  lateral  lobes  a  large  and  distinct  spiracle,  in  the  cast  skin  with  the 
large  tracheal  trunks  still  attached.  Female  cauda  (Plate  XXXV,  156)  stout,  tergal  valves 
a  little  longer  than  sternal  valves  and  a  little  upcurved. 

Nepionotype.—  Ithaca,  New  York,  October  14,  1912. 

Neanotype. —  With  type  larva. 

Paratypes.—  Type  locality,  October  12  to  19,  1912.    , 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  843 

Subtribe  Epiphragmaria 

So  far  as  is  known  to  the  writer,  the  division  Epiphragmaria  includes 
only  the  genus  Epiphragma.  The  immature  stages  are  of  a  very  primitive 
organization.  The  larvae  have  the  head  capsule  broad  and  massive, 
with  the  posterior  incisions  very  shallow.  The  labrum  is  broad,  with 
lateral  tufts  of  hair.  The  mentum  is  heavily  chitinized,  entire,  with 
only  a  single  lateral  tooth  on  either  side  of  the  median  tooth.  The 
hypopharynx  is  not  chitinized  but  the  entire  surface  is  provided  with 
rows  of  tiny  spinous  points,  somewhat  as  in  the  Dicranoptycharia.  The 
antennae  have  the  terminal  papillae  hemispherical.  The  mandibles  are 
not  greatly  elongated;  each  has  an  apical  tooth  and  two  rows  of  lateral 
teeth.  The  maxillae  are  very  simple  in  structure.  The  abdomen  is 
provided  with  six  ventral  creeping-welts.  The  spiracular  disk  is  large 
and  simple,  with  the  lobes  short  and  obtuse. 

The  pupa  has  the  breathing  horns  heavily  chitinized,  tapering  to  acute, 
slender  points.  The  spiracles  on  the  dorsum  of  the  eighth  abdominal 
segment  are  large  and  conspicuous. 

The  relationships  of  this  group  are  obviously  with  the  Rhamphidaria  of 
the  Limnobiini  and  the  Ularia  of  the  Hexatomini,  the  three  groups  probably 
being  closer  together  phylogenetically  than  their  arrangement  in  tribes 
would  indicate.  From  a  study  of  the  adults  alone,  there  seems  to  be 
little  connection  between  the  genera  Rhamphidia,  Epiphragma,  and  Ula, 
which  were  placed  in  three  widely  separated  tribes  by  Osten  Sacken.  A 
study  of  the  immature  stages,  the  structure  of  the  larval  head,  the  abdom- 
inal creeping- welts,  and  the  dorsal  spiracles  of  the  eighth  abdominal 
segment  of  the  pupa,  indicate  a  relationship  that  cannot  be  denied. 

Genus  Epiphragma  Osten  Sacken  (Gr.  upon  +  partition) 
1859    Epiphragma  0.  S.    Proc.  Acad.  Nat.  Sci.  Phila.*  p.  238. 

Larva.—  Body  nearly  smooth;  basal  annul!  of  abdominal  segments  2  to  7  with  a  naked 
transverse  creeping-welt  on  ventral  surface.  Spiracular  disk  large,  with  four  or  five  lobes, 
the  dorso-median  lobe  often  reduced.  Spiracles  large,  circular.  Anal  gills  four,  retractile. 
Head  capsule  massive.  Labrum  broad,  with  lateral  tufts  of  hairs  and  with  two  setae  on 
anterior  margin.  Mandible  with  a  blunt  apical  tooth  and  two  rows  of  two  lateral  teeth. 
Maxilla  small,  simple;  palpus  large;  inner  lobe  with  a  dense  tuft  of  stiff  hairs.  Antenna 
two-segmented;  basal  segment  elongate,  apical  papilla  hemispherical.  Hypopharynx  not 
chitinized.  Mentum  entire,  with  only  three  teeth  —  a  slender  median  tooth,  and  a  broad, 
fat  tooth  on  either  side. 


844  CHARLES  PAUL  ALEXANDER 


.^  Cephalic  crest  erect,  rectangular,  the  outer  lateral  angles  produced  into  strong, 
chitinized  spines.  Pronotal  breathing  horns  large  at  base,  the  elongate  tips  nearly  chitinized, 
tapering  into  subacute  points.  Wing  sheaths  ending  opposite  base  of  third  abdominal 
segment.  Leg  sheaths  terminating  just  before  end  of  fifth  abdominal  segment.  Abdominal 
tergites  with  a  transverse  band  of  spicules  at  caudal  margin;  pleurites  with  a  transverse 
oval  area  of  similar  spines;  terminal  sternites  with  a  similar  transverse  band  of  spicules, 
most  powerfully  developed  on  segment  8;  dorsal  spiracles  on  segment  8  conspicuous. 

Epiphragma  is  a  rather  small  genus  (twenty-five  species)  of  usually 
large  and  handsome  crane-flies,  finding  its  center  of  distribution  in  the 
tropical  regions  of  the  New  World.  The  adult  flies  of  the  local  species 
are  usually  common.  Epiphragma  fascipennis,  the  best  known  of  these 
species,  is  often  found  swarming  in  shady  places,  the  swarms  usually 
consisting  of  from  twelve  to  fifteen  individuals. 

The  immature  stages  of  all  the  species  thus  far  known  are  spent  in  decay- 
ing wood.  In  Europe,  Beling  (  1  873  b:  589-590)  records  finding  larvae  and 
pupae  of  E.  picta  (Fabr.)  in  decaying  ash  (Fraxinus)  and  beech  (Fagus) 
in  the  spring.  He  gives  the  pupal  duration  of  this  species  as  about  two 
weeks.  Needham  (1903:281-285)  gives  an  excellent  account  of  E. 
fascipennis,  which  he  found  in  partly  decayed  stems  of  willow  and  button- 
bush.  The  same  species  has  been  found  living  in  the  stems  of  black 
ash  and  elm.  E.  solatrix  has  been  found  in  various  deciduous  trees  in 
a  dead  or  decaying  condition.  Bruch  (in  litt.)  records  an  Argentinian 
Epiphragma  from  decaying  wood. 

Epiphragma  solatrix  (0.  S.) 

1859    Limnophila  (Epiphragma)  solatrix  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  238. 

Epiphragma  solatrix  is  a  beautiful  crane-fly,  a  little  more  Austral  in 
its  distribution  than  the  more  widely  distributed  E.  fascipennis.  H.  S. 
Barber  found  larvae  and  young  pupae  very  numerous  in  a  drift  log  of 
sycamore  (Platanus)  at  Plummers  Island,  Maryland,  on  May  19,  1913, 
and  it  is  the  study  of  this  material  that  is  included  in  this  paper.  Shannon 
found  larvae  in  rotten  wood  at  Rosslyn,  Virginia,  on  November  23  and 
25,  1912,  and  a  pupa  on  January  3,  1913.  Later  he  reared  both  E. 
solatrix  and  E.  fascipennis  from  larvae  taken  from  the  same  log  on  the 
same  day.  The  species  was  'again  found  in  a  decaying  maple  log  on  the 
Potomac  River  opposite  Plummers  Island,  Maryland,  where  it  was  associ- 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  845 

ated  with  the  larvae  of  the  syrphid  fly,   Temnostoma  bombylans  (Fabr.) 
and  the  supposed  larva  of  Protoplasa  (p.  770). 

Larva. —  Length,  18  mm. 

Diameter,  1.6  mm. 

Coloration  pale  whitish;  anterior  segments  of  body  brighter-colored,  more  reddish. 

Form  stout  and  plump,  not  narrowed  at  ends  of  body.  Pronotum  longer  than  other 
thoracic  segments.  Integument  covered  with  a  short  appressed  pubescence.  Abdominal 
segments  2  to  7  with  conspicuous  white  creeping-welts  on  ventral  surface  of  basal  ring;  these 
welts  not  interrupted  medially,  and  entirely  naked.  Spiracular  disk  (Plate  XXXVI,  163) 
large,  unmarked,  surrounded  by  five  short  lobes,  the  ventral  pair  very  widely  separated; 
dorsal  lobe  broad  and  obtuse;  entire  disk  fringed  with  very  short,  delicate  hairs  which  are 
interrupted  only  between  paired  lobes;  inner  face  of  lobes  suffused  with  pale  brown  at  tips. 
Spiracles  large,  circular,  reddish  with  a  very  broad  yellow  margin,  separated  by  a  distance 
nearly  equal  to  two  times  the  diameter  of  one  spiracle.  Anal  gills  four,  retractile. 

Head  capsule  (Plate  XXXVI,  158)  broad  and  massive,  the  constituent  plates  firmly  united 
except  on  ventral  parts.  Labrum  broadly  transverse;  a  median  epipharyngeal  part  whose 
surface  is  finely  pitted,  bearing  on  ventral  side  near  margin  two  stout  setae;  lateral  angles  of 
labrum  directed  proximad  and  densely  tufted  with  hairs;  extreme  lateral  margins  of  labrum 
with  a  curved  hyaline  seta;  juncture  of  labrum  and  clypeus  with  two  large  setiferous  punctures 
on  either  sid3;  a  few  setiferous  punctures  on  head  capsule.  Mentum  (Plate  XXXVI,  159) 
heavily  chitinized,  the  outer  face  terminating  in  a  single  median  tooth,  behind  which  is  a 
tridentate  plate  with  three  flattened  teeth,  the  lateral  ones  very  broad.  Prementum  (Plate 
XXXVI,  160)  lying  just  behind  men  turn,  a  moderately  broad  transverse  plate  who£e 
anterior  margin  is  deeply  and  almost  squarely  notched,  the  lateral  lobes  thus  formed  being 
rounded  or  feebly  indented  at  their  tips.  Hypopharynx  (Plate  XXXVI,  161)  lying  above 
and  connected  with  prementum,  consisting  of  a  large  flattened  lobe,  whose  outer  margin 
is  evenly  rounded,  the  surface  densely  set  with  longitudinal  rows  of  small,  subr.cute  papillae. 
Antenna  short,  two-segmented;  basal  segment  cylindrical,  with  auditory  plate  almost  basal 
in  position;  apical  segment  small,  subglobular  or  hemispherical.  Mandible  moderate  in 
size,  broad  at  base  with  a  blunt  apical  tooth  and  a  few  blunt  lateral  teeth,  two  on  dorsal 
cutting  edge  and  two  on  ventral  cutting  edge,  the  more  basal  of  these  latter  tending  to  be 
evanescent;  two  stiff  setae  at  heel  of  mandible;  a  triangular  lobe  on  dorsal  face  which 
projects  onto  lateral  parts  of  labrum.  Maxilla  (Plate  XXXVI,  162)  primitive  in  structure; 
cardo  elongate,  transverse,  with  four  setiferous  punctures  bearing  powerful  setae;  outer  lobe 
glabrous,  with  palpus  at  its  tip;  palpus  slightly  elongate,  the  auditory  plate  just  before 
midlength;  a  seta  on  outer  lobe  below  palpus;  inner  lobe  of  maxilla  with  a  dense  terminal 
brush  of  stiff  yellow  hairs  and  a  few  sensory  papillae. 

Pupa. —  Length,  12-12.5  mm. 

Width,  d.-s.,  2-2.1  mm. 
Depth,  d.-v.,  2-2.1  mm. 

Head,  thorax,  and  appendages  brown,  becoming  darker  with  age;  pronotal  breathing 
horns  light  red;  abdomen  whitish,  the  terminal  rows  of  spicules  on  the  segments  chitinized. 


846  CHARLES  PAUL  ALEXANDER 

(In  older  specimens  the  wing  pattern  shows  on  the  sheath  and  is  not  distinctly  fasciate  as 
in  E.  fascipennis.) 

Cephalic  crest  (Plate  XXXVII,  166)  erect,  rectangular,  the  outer  lateral  angles  directed 
ventrad  and  laterad  as  powerful  chitinized  spines,  which  in  most  cases  curve  outward;  a 
stout  seta  beneath  each  spine.  Labrum  subtruncated  or  indistinctly  bilobed  at  apex. 
Labial  lobes  triangular,  projecting  caudad  as  two  cones.  Maxillary  palpi  bent  backward, 
lying  along  margin  of  cheek,  stout,  tapering  suddenly  to  blunt  tips.  Cheeks  projecting 
as  flattened  ledges.  Antennae  moderate  in  length,  on  scapal  segment  with  a  flattened 
tubercle  on  either  side  of  cephalic  crest;  antennae  extending  rather  far  beyond  wing 
root. 

Pronotal  breathing  horns  (Plate  XXXVII,  164)  with  base  enlarged,  the  elongate,  subchiti- 
nized  tips  gradually  narrowed,  and  the  extreme  apices  subacute;  breathing  horns  at  tips 
cirected  ventrad  and  proximad;  a  truncated  setiferous  tubercle  before  each  breathing  horn. 
Mesonotum  very  deep.  A  short,  but  high,  median  crest  behind  breathing  horns.  Lateral 
angles  of  thorax  produced  into  a  slender  lobe  bearing  a  stout  seta  at  tip.  Wing  sheaths 
ending  opposite  base  of  third  abdominal  segment.  Leg  sheaths  terminating  just  before  end 
of  fifth  abdominal  segment;  middle  tarsal  sheaths  usually  a  little  shorter  than  the  others;  in 
some  specimens,  especially  females,  fore  legs  the  shortest,  hind  legs  the  longest.  Abdominal 
segments  (Plate  XXXVII,  167)  with  two  narrow  basal  rings  and  a  broader  posterior  ring. 
Tergites  on  segments  2  to  7  near  caudal  margin  of  posterior  ring  with  a  narrow  transverse 
crossband  densely  beset  with  acute  spicules;  on  posterior  segments  these  bands  becoming 
gradually  weaker,  on  segment  7  being  very  weak;  two  setae  near  lateral  margin  of  posterior 
ring  at  about  midlength.  Pleurites  with  a  transverse  oval  area  with  about  twenty  sharp  spines 
near  caudal  margins  of  segments;  a  stout  seta  lying  cephalad  of  this  area.  Spiracles  distinct, 
oval,  lying  on  ventral  cephalic  part  of  pleural  posterior  ring;  a  seta  near  dorsal  margin  of 
second  basal  ring.  Sternites  with  a  transverse  band  of  spicules  on  segments  5  to  8,  these 
being  weakest  on  segment  5  at  ends  of  tarsal  sheaths  and  very  strong  and  powerful  on  seg- 
ment 8,  where  the  band  is  interrupted  medially;  band  often  rather  narrow  but  sometimes 
much  broader.  A  seta  near  midlength  of  posterior  ring  on  either  side.  Male  cauda  (Plate 
XXXVIII,  170)  with  dorsal  lobes  very  stout,  globose  at  base  but  soon  narrowed  into  slender 
cylindrical  appendages,  each  bearing  three  setae  at  tip;  viewed  from  above,  these  lobes  are 
seen  to  be  separated  by  a  wide  U-shaped  notch;  ventral  lobes  blunt  and  with  a  flattened 
depressed  disk  at  base  of  incision;  eighth  segment  with  a  wide  rectangle  of  lobes,  two 
posterior  lobes  that  are  weakly  setiferous  and  two  smaller  anterior  lobes  with  a  large 
spiracle  proximad  of  each.  Female  cauda  (Plate  XXXVIII,  171)  with  tergal  valves  a  little 
longer  than  sternal  valves,  upcurved,  and  with  a  stout  seta  before  tip. 

Nepionotype. —  Plummers  Island,  Maryland,  May  19,  1913. 

Neanotype. —  With  type. 

Paratypes. —  Larvae  and  pupae  with  type. 

Epiphragma  fascipennis  (Say) 

1823    Limnobia  fascipennis  Say.     Journ.  Acad.  Nat.  Sci.  Phila.,  vol.  3,  p.  19. 

1859    Limnophila  (Epiphragma)  pavonina  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  239. 

1869     Epiphragma  fascipennis  O.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  194. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  847 

Epiphragma  fascipennis  is  a  very  common  crane-fly  thruout  eastern 
North  America.  As  already  stated,  the  immature  stages  are  spent  in 
decaying  or  partly  sound  wood,  a  wide  variety  of  deciduous  trees  and 
shrubs  being  chosen,  such  as  willow,  elm,  ash,  buttonbush,  and  others. 
Needham  gives  the  pupal  duration  as  about  twelve  days,  larvae  and 
pupae  found  on  May  18  emerging  as  adults  on  the  30th.  A  fully  grown 
larva  that  the  writer  found  in  a  decayed  log  beneath  moss  at  Ithaca, 
New  York,  on  May  8,  1917,  pupated  early  in  the  morning  of  the  10th. 
The  specimen  died  on  the  18th,  when  about  to  emerge,  and  this  would 
give  a  much  shorter  pupal  period  than  is  generally  recorded  for  the  genus. 
It  was  noted  at  the  same  time  that  the  larva  superficially  resembles  the 
larva  of  the  leptid  fly  Chrysopila  thoracica  (Fabr.),  with  which  it  was 
associated  but  from  which  it  is  easily  distinguished  by  its  massive  he*ad 
capsule. 

Needham  found  abundant  pupae  in  a  decaying  log  of  black  ash  (Fraxinus) 
near  Freeville,  New  York,  on  May  6,  1915.  The  pupae  occurred  in  bur- 
rows in  the  semi-decayed  wood.  Adults  emerged  on  the  llth.  Additional 
material  was  found  at  Mud  Creek,  near  Freeville,  on  May  15,  1915,  in 
elm  (Ulmus). 

The  account  of  the  habits  of  the  larvae  as  observed  in  Illinois  by 
Needham  (1903:281-285)  is  here  quoted  in  part: 

The  larvae  bore  in  the  dead  and  fallen  stems  of  buttonbush  and  willow,  where  these  lie 
on  the  mud  at  the  borders  of  shallow  ponds.  I  found  them  always  in  stems  that  were  still 
partially  sound,  tunneling  beneath  the  bark  or  even  into  the  deeper  parts  and  into  the 
sounder  wood.  These  stems  are  frequently  submerged  in  spring  and  autumn,  and  even 
in  summer,  when  the  pond  has  gone  dry,  they  are  always  saturated  with  moisture 

The  most  interesting  thing  about  the  larva,  aside  from  its  wood-boring  habits,  is  its  singular 
adaptation  to  amphibian  life.  It  must  needs  live  part  of  the  time  wholly  submerged  beneath 
the  waters  of  the  pond,  and  part  of  the  time  out  on  land;  it  has,  therefore,  both  open  spiracles 
and  tracheal  gills;  and,  moreover,  its  tracheal  gills  are  so  placed  that  they  may  be  with- 
drawn into  the  body  in  a  dry  time,  where  they  escape  the  ills  of  too  rapid  evaporation. 

In  his  description  of  the  immature  stages,  Needham  points  out  a 
probable  error  of  Beling  in  describing  a  sexual  dimorphism  in  the  larvae 
of  a  species  of  this  genus  —  Beling  stating  that  the  larvae  producing 
females  have  three  caudal  lobes  while  those  producing  males  have  five. 
Malloch  (1915-17  b:  224-225)  cites  Needham' s  descriptions  of  this  species. 

Larva.- —  (No  larvae  are  available  to  the  writer  for  a  comparison  with  this  stage  of 
Epiphragma  solatrix,  but  from  Needham's  characterization,  and  manuscript  notes  on  speci- 
mens taker  at  Ithaca,  New  York,  by  the  writer,  the  following  differences  seem  to  hold) : 


848  CHARLES  PAUL  ALEXANDER 

Spiracular  disk  surrounded  by  but  four  lobes,  the  dorso-median  lobe  being  very  reduced; 
inner  faces  of  these  lobes,  as  well  as  disk  itself,  entirely  unmarked  with  darker. 
Pupa.- — Similar  to  pupa  of  Epiphragma  solatrix,  but  differing  as  follows: 
Cephalic  crest  (Plate  XXXVII,  165)  low,   the  lateral  horns  shorter,   not  elevated,  and 
directed  ventrad   or   slightly   downward;   setae   on   lateral   face  of   spines  often  projecting 
above  (cephalad  of)   it.     Pronotal  breathing  horns  shorter  and   stouter  than   in   solatrix. 
Spines  on  eighth  abdominal  sternite  widely  separated  or  interrupted  on  median  line.     Male 
cauda  (Plate  XXXVIII,  169)  with  dorsal  lobes  much  stouter  than  in  solatrix. 

Neanotype.—  Bool's  hillside,  Ithaca,  New  York,  May  8,  1917. 
Paratype.—  Pupa,  Mud  Creek,  Freeville,  New  York,  May  15,  1915. 

The  type  pupa  has  one  of  the  pronotal  breathing  horns  deformed  and 
bent  caudad  so  as  to  be  appressed  against  the  body.  Needham,  who 
has  a  very  much  larger  series  available  for  study,  notes  this  same  peculiarity 
when  he  states  that  "a  crumpled  horn  on  one  side  is  of  rather  common 
occurrence"  (Needham,  1903:284). 

Subtribe  Pseudolimnophilaria 

As  at  present  constituted,  the  division  Pseudolimnophilaria  includes 
the  single  genus  Pseudolimnophila,  but  other  groups  may  be  added  to 
it  when  the  immature  stages  of  other  species  of  the  old  genus  Limnophila 
are  made  known.  The  larva  is  of  a  distinctly  primitive  type,  the  head 
capsule  being  compact  and  massive,  the  mentum  chitinized  and  com- 
pletely divided,  each  half  provided  with  seven  or  eight  teeth.  The 
hypopharynx  is  a  heavily  chitinized  semicircle  with  numerous  teeth  around 
the  anterior  margin.  The  antennae  bear  two  elongate  apical  papillae. 
The  mandibles  each  have  two  blunt  apical  teeth  and  two  rows  of  lateral 
teeth. 

The  pupa  is  similar  to  that  of  the  Limnophilaria,  but  the  abdominal 
segments  have  transverse  rows  of  setiferous  tubercles. 

The  larva  shows  many  points  of  resemblance  to  the  tribe  Limnobiini, 
and  the  two  groups  are  probably  derivable  from  an  immediate  common 
ancestor. 

Genus  Pseudolimnophila  Alexander  (Gr.  false  +  Limnophila) 

1919     Pseudolimnophila  Alex.     Cornell  Univ.  Agr.  Exp.  Sta.,  Mem.  25,  p.  917. 

Larva. —  Body  covered  with  delicate  appressed  hairs  and  numerous  erect  setae.  Spiracular 
disk  surrounded  by  four  lobes,  the  ventral  pair  very  long  and  narrow,  with  long  fringes  of 
hair.  Gills  four,  long  and  slender.  Head  capsule  massive,  the  sclerites  large,  fused. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  849 

Mandible  short  and  blunt,  of  the  generalized  limnobiine  type,  the  two  apical  teeth  blunt. 
Maxilla  not  greatly  projecting.  Antenna  slender,  with  two  elongate  apical  papillae.  Hypo- 
pharynx  a  chitinized  band,  with  numerous  teeth  along  anterior  margin.  M^ntum  completely 
divided,  each  half  with  about  seven  teeth. 

Pupa. —  Cephalic  crest  tipped  with  long  setae.  Pronotal  breathing  horns  elongate, 
tips  deeply  split  into  divergent  flaps.  Abdominal  segments  with  transverse  rows  of  setiferous 
tubercles,  there  being  five  such  rows  on  the  tergites  and  six  on  the  sternites.  Lateral  spiracles 
protuberant. 

The  name  Pseudolimnophila  was  proposed  for  a  certain  group  of  the 
old  genus  Limnophila,  including  L.  luteipennis  and  its  allies.  The  adult 
flies  closely  resemble  species  of  Limnophila,  but  the  immature  stages  are 
very  different  and  of  a  distinctly  more  generalized  type. 

The  larvae  are  herbivores,  with  the  mentum  heavily  chitinized,  com- 
pletely divided  into  two  halves,  the  margin  conspicuously  toothed,  the 
hypopharynx  heavily  chitinized  and  with  numerous  teeth,  and  the  mandi- 
bles short  and  blunt  and  with  many  obtuse  teeth.  The  points  of  resem- 
blance to  the  Limnobia  type  are  numerous,  and  the  head  capsule  shows 
few  features  in  common  with  Limnophila,  in  the  restricted  sense.  The 
pupae  have  the  abdomen  with  numerous  transverse  rows  of  setiferous 
tubercles  bearing  long  setae.  The  genotype  is  Limnophila  luteipennis 
O.  S.,  of  eastern  North  America.  Other  species  referable  to  this  genus 
are  L.  inornata,  L.  contempta,  L.  nigripleura,  and  L.  noveboracensis,  of  the 
Nearctic  fauna;  L.  lucorum  (Meig.),  of  Europe;  and  L.  frugi  Bergr., 
L.  claduroneura  Speis.,  L.  natalensis  Alex.,  L.  spectabilis  Alex.,  and  several 
other  species,  of  the  Ethiopian  fauna. 

Hart  (1898  [18951:202-204)  describes  the  habits  of  P.  luteipennis  in 
some  detail.  Larvae  about  half  grown  were  found  on  March  17,  1895, 
these  attaining  their  growth  and  pupating  on  April  13.  Hart  ascertained 
by  dissection  that  the  species  is  a  herbivore,  feeding  on  fragments  of 
dead  vegetation,  numerous  diatoms  of  many  species,  and  minute  algae. 

As  in  many  species  of  the  Limnophilaria  and  a  few  other  groups  of 
crane-flies,  the  spiracular  lobes  are  fringed  with  long,  delicate  hairs  which 
spread  out  on  the  surface  film  of  the  water  into  broad  fans.  The  larva 
is  unable  to  release  itself  from  the  hold  of  the  water  except  by  looping 
the  head  and  the  anterior  end  of  the  body  around  the  posterior  end,  and 
drawing  the  latter  thru  this  loop,  repeating  this  action  until  the  hairs 
have  gone  below  the  level  of  the  film. 
6 


850  CHARLES  PAUL  ALEXANDER 

It  is  probable  that  other  groups  of  species  of  the  old  genus  Limnophila 
will  be  found  to  deviate  from  the  characters  of  this  group  as  now  restricted. 
Species  such  as  L.  hyalipennis  (Zett.),  L.  nemoralis  (Meig.),  and  others  in 
Europe,  and  L.  brevifurca,  L.  emmelina,  L.  mundoides,  L.  toxoneura, 
L.  ultima,  and  similar  anomalous  forms  in  America,  should  be  investigated 
critically  when  opportunity  offers. 

Pseudolimnophila  luteipennis  (0.  S.) 

1859    Limnophila  luteipennis  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  236. 

Pseudolimnophila  luteipennis  is  one  of  the  most  abundant  species  of 
the  tribe,  and  is  very  widely  distributed  thruout  eastern  North  America. 
The  adult  flies  are  common  in  swamps  and  along  the  marshy  edges  of 
streams  and  ponds.  The  immature  stages  are  characteristic  swamp 
inhabitants.  They  have  been  described  in  detail  by  Hart  (1898  [1895] : 
202-204).  Malloch  (1915-17  b :  222-223)  adds  several  supplementary  notes 
and  figures  of  Hart's  material. 

Larva. —  Length,  15-18  mm. 

Diameter,  1.5-1.7  mm. 

Coloration  light  brownish  yellow,  slightly  darker  toward  posterior  end  of  body. 

Body  covered  with  a  delicate  appressed  brownish  pubescence  and  abundant  stiff  bristles 
and  erect  hairs  on  each  segment,  arranged  in  about  five  transverse  rows;  these  bristles  more 
numerous,  coarser,  and  blacker,  on  posterior  segments,  forming  a  large  tuft  on  sides  of 
penultimate  segment  of  body.  Spiracular  disk  (Plate  XXXIX,  176)  surrounded  by  four 
lobes,  the  posterior  pair  very  long  and  slender,  finger-like,  the  lateral  pair  much  shorter  but 
slender;  the  inner  face  of  these  lobes  with  brownish  black  or  dark  brown  transverse  lines  which 
cover  almost  the  entire  face;  an  arcuate  line  extending  from  ends  of  ventral  marks  across  disk 
between  spiracles;  two  small  triangular  marks  located  between  ventral  lobes;  lobes  fringed 
with,  long,  delicate  hairs,  those  at  ends  very  elongate,  those  toward  base  of  lobes  shortened 
but  continuous  around  disk.  Anal  gills  four,  very  long  and  slender,  exceeding  ventral  lobes 
of  spiracular  disk. 

Head  capsule  short  and  stout,  exhibiting  a  very  generalized  condition.  Prefrons  very 
broad,  only  slightly  narrowed  behind  and  but  slightly  exceeding  lateral  plates.  Labrum 
broadly  transverse,  with  a  large  tuft  of  hairs  on  either  side.  Mentum  (Plate  XXXIX,  172) 
completely  divided,  each  half  with  seven  (or  rarely  eight)  teeth  along  anterior  margin,  of 
which  the  third  (or  fourth)  from  the  inside  is  the  largest;  a  flattened  lobe  just  laterad  of  each 
half  of  mentum;  mental  plates  continuous  behind  with  strong  lateral  plates  of  capsule;  in 
a  normal  position  the  two  halves  of  mentum  overlying  each  other  to  a  greater  or  less  degree. 
Hypopharynx  (Plate  XXXIX,  173)  with  anterior  margin  broadly  roundSd,  with  numerous 
(from  ten  to  fifteen)  bluntly  rounded  teeth.  Antenna  slender,  with  two  long  apical 
papillae.  Mandible  (Plate  XXXIX,  175)  short  and  broad,  of  the  generalized  limnobiine 


THE  CRANE-FLIES  or  NEW  YORK  —  PART  II  851 

type;  tips  blunt,  ending  in  two  large  approximated  teeth,  with  a  row  of  smaller  lateral  teeth 
extending  basad  of  each;  ventral  cutting  edge  with  about  four  such  lateral  teeth,  dorsal  edge 
with  one  or  two  much  larger  teeth;  two  strong  setae  near  base  of  scrobe,  and  two  others  near 
tip  of  mandible;  a  large  brush  of  hairs  at  prosthecal  region.  Maxilla  not  projecting,  with 
dense  brushes  of  short,  stiff  hairs  and  with  two  short  sensory  tubercles  which  are  tipped  with 
very  elongate  setae;  palpi  stout  and  chitinized  at  base,  the  apex  narrowed  and  pale. 

Pupa. —  Length,  10-13  mm. 

Width,  d.-s.,  about  1.5  mm. 

Thorax  reddish  brown  to  black  in  color,  depending  on  age  of  specimen;  abdomen  dirty 
whitish  with  narrow  transverse  brownish  lines,  both  above  and  below;  pronotal  breathing 
horns  dark  brown  or  blackish. 

Form  subcylindrical,  abdomen  slightly  depressed.  Cephalic  crest  of  two  slender  tubercles 
behind,  tipped  with  long  setae;  another  pair  of  setae  between  antennal  bases.  Pronotal 
breathing  horns  elongate,  transversely  ringed,  the  tips  widely  and  deeply  split  into  two 
divergent  flaps.  Prothorax  with  a  prominent  median  carina.  A  tubercle  in  front  of  each 
breathing  horn,  each  with  about  four  stout  setae.  Mesonotum  above  wing  axil  with  a 
tubercle  bearing  throe  setae.  Wings  reaching  end  of  second  abdominal  segment.  Legs 
reaching  end  of  third  abdominal  segment,  the  tarsi  ending  about  on  a  level  or  those  of  fore 
legs  a  little  longer  than  those  of  other  two  pairs. 

Abdominal  segments  divided  into  two  annuli,  the  basal  one  still  further  subdivided  into 
annulets;  on  tergum  three  annulets,  each  bearing  a  transverse  row  of  setiferous  tubercles,  the 
setae  very  long  and  conspicuous,  in  some  cases  the  tubercles  multisetose,  third  annulet 
with  two  widely  separated,  slender,  elongate  tubercles.  Posterior  annulus  with  a  basal 
and  a  terminal  transverse 'row  of  setiferous  tubercles.  On  sternum,  four  transverse  rows  of 
setiferous  tubercles  on  basal  ring  and  two  on  posterior  ring.  On  pleura,  protuberant  spiracles. 
Lateral  angles  of  segment  8  jutting  out  into  stout  lobes  which  are  densely  studded  with 
setiferous  tubercles,  on  ventral  face  continued  toward  median  line  as  a  nearly  straight  row 
of  about  eight  setiferous  tubercles,  on  dorsum  a  finger-like  lobe  on  either  side:  Male  cauda 
(Plate  XL,  178)  with  sternal  valves  shorter  than  dorsal  lobes,  blunt  at  tips;  tergal  valves 
acutely  pointed,  directed  dorsad.  Female  cauda  (Plate  XL,  179  and  180)  with  tergal  valves 
considerably  longer  than  the  more  slender  sternal  valves,  broad  medially,  tapering  to  the 
broad,  blunt  tips  which  terminate  in  blackened  points;  two  setae  on  outer  margin. 

Nepionotype. —  Larch  Meadows,  Ithaca,  New  York,  April  20,  1917. 

Neanotype. —  Female  pupa,  with  type.     No.  19-1917. 

Paratypes. —  Larvae  with  type.      Pupae  of  both  sexes  from  Bool's  hillside,  Ithaca,  New 

York,  June  6,  1917.     A  female  pupa,  Orono,  Maine,  taken  as  a  larva  June  13,  1913, 

emerged  June  22.     Female  pupa,  July  15,  1913,  No.  75-1913. 

Pseudolimnophila  inornata  (0.  S.) 

1869    lAmnophila  inornata  0.  S.    Mon.  Dipt.  N.  Amer.,  part  4,  p.  219,  220. 
Pseudolimnophila  inornata  is  not  so  common  as  P.  luteipennis  but  is 
found  in  similar  situations.     The  immature  stages  are  spent  in  rich  organic 
mud.     The  only  specimens  that  the  writer  has  reared  were  taken  in 


852  CHARLES  PAUL  ALEXANDER 

Larch  Meadows,  near  Ithaca,  New  York,  on  May  15,  1917,  where  they 
were  associated  with  larvae  of  Rhamphidia  mainensis,  Pseudolimnophila 
luteipennis,  Tipula  dejecta,  and  other  swamp-inhabiting  species. 

Larva.- —  Very  similar  to  that  of  P.  luteipennis,  but  body  is  darker  and  is  conspicuously 
blotched  with  whitish,  especially  on  posterior  parts  of  ventral  segments.  Mouth  parts 
nearly  the  same  in  the  two  species.  Antenna  (Plate  XXXIX,  174)  with  two  very  long  and 
slender  terminal  papillae,  one  blunt  at  tip,  the  other  much  longer  and  tapering  gradually 
to  tip. 

Pupa.- —  Very  similar  to  that  of  P.  luteipennis,  but  somewhat  smaller  and  darker-colored. 
Pronotal  breathing  horns  (Plate  XL,  177)  darker  brown,  the  divergent  terminal  flaps  pro- 
portionately a  little»more  elongate,  equal  to  almost  one-quarter  length  of  entire  horn.  Dorsal 
lobes  at  base  of  ovipositor  more  attenuated. 

Nepionotype  —  Larch  Meadows,  Ithaca,  New  York,  May  15,  1917.     No.  53-1917. 
Neanotype  —  Larch  Meadows,  Ithaca,  New  York,  May   25,    1917.      No.  53-1917,    cast 
pupal  skin. 

Subtribe  Dactylolabaria 

The  division  Dactylolabaria  is  proposed  for  the  genus  Dactylolabis, 
a  small  group  of  curious  crane-flies  which  are  still  not  well  understood. 
The  adult  flies  bear  a  striking  resemblance  to  species  of  the  genus  Lim- 
nophila,  but  the  larvae  and  the  pupae  show  characters  that  are  not  found 
in  the  more  specialized  divisions  of  the  Hexatomini.  The  type  of  the 
genus  is  Limnophila  montana  O.  S.  of  the  eastern  United  States.  Other 
Nearctic  species  included  are  L.  damula  O.  S.  (western  United  States), 
L.  rhicnoptiloides  Alex,  (northwestern  Canada),  L.  cubitalis  0.  S.  (eastern 
United  States),  L.  nitidithorax  Alex,  (western  United  States),  and  L. 
hortensia  Alex,  (western  Canada).  ,  \ 

The  immature  stages  of  two  European  species  —  D.  wodzickii  (Now.) 
and  D.  denticulata  Bergr. —  have  been  made  known  by  Nowicki  (1867: 
340-343)  and  by  Mik  (1894),  respectively.  Their  descriptions  of  the 
details  of  the  larval  head  anxl  the  lateral  spiracles  of  the  abdomen  of  the 
pupa  are  incomplete,  however.  The  larvae  of  none  of  the  American 
species  have  as  yet  been  made  known,  but  the  pupae  of  D.  cubitalis  O.  S. 
were  found  by  Needham  and  are  described  hereinafter. 

Genus  Dactylolabis  Osten  Sacken  (Gr.  finger     forceps) 
1859    Dactylolabis  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  240. 

Larva. —  Body  very  depressed,  the  ventral  surface,  especially,  being  greatly  flattened. 
Head  capsule  compact;  mentum  not  completely  divided,  its  anterior  margin  with  seven 
teeth.  Mandible  with  but  few  lateral  teeth. 


I 

THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  853 

Pupa.—  Somewhat  similar  to  pupae  of  Limnophilaria.  Pronotal  breathing  horns  short 
and  cylindrical.  In  some  species,  at  least,  second  abdominal  segment  with  a  very  large,  and 
prominent  lateral  spiracle. 

Dactylolabis  is  a  small  genus  including  about  thirteen  known  species 
which  are  about  equally  well  distributed  in  Europe  and  North  America. 

Dactylolabis  denticulata  (Bergr.) 

1891    Limnophila  denticulata  Bergr.    Mittheil.  Naturf.  Ges.  Bern,  p.  132. 

The  present  knowledge  of  the  life  history  of  Dactylolabis  denticulata 
is  due  to  the  work  of  Mik  (1894),  whose  paper  is  cited  by  other  workers, 
among  them  Grtinberg  (1910:54-55). 

Larvae  and  pupae  were  found  on  the  dark  brownish  gray  chalk  cliffs 
near  Steiermark  on  August  3,  1891.  In  certain  places  on  the  cliffs  were 
broad  or  narrow  bands  of  black.  The  larvae  and  the  pupae  were  found 
along  these  dark  bands  and  showed  a  decided  resemblance  to  their  sur- 
roundings. The  surface  of  the  rock  was  wet  with  dripping  water,  which 
supported  a  flora  of  lowly  plant  organisms  on  which  the  larvae  presumably 
fed.  The  adherence  of  the  pupa  to  the  last  larval  skin  is  of  interest  since 
it  presumably  aids  in  the  emergence  of  the  adult.  A  similar  condition 
is  found  in  Cylindrotoma. 

Larva. —  Length,  7  mm. 
Width,  2  mm. 

Form  very  depressed  (Plate  XLI,  183),  the  ventral  surface  flattened,  almost  leechlike. 
Both  dorsal  and  ventral  surface  provided  with  hairs,  with  longer  and  more  numerous  hairs 
on  sides  of  body.  Body  covered  with  small,  black,  structureless  particles  of  earth  or  excre- 
ment, more  numerous  near  margins  of  dorsal  surface,  these  particles  lacking  a  definite  arrange- 
ment and  being  very  firmly  attached  to  the  body  hairs;  when  particles  are  removed,  larva 
is  of  a  greenish  gray  color,  subhyaline.  Abdominal  segments  divided  into  two  rings  bearing 
long  hairs  which  are  more  conspicuous  on  sides  of  body.  Head  capsule  (Plate  XLI,  182) 
small  but  compact,  completely  retractile  within  prothoracic  segment.  Labrum  relatively 
large  and  semitransparent,  the  margins  with  long  hairs;  on  dorsum  on  either  side  a  long 
delicate  seta.  Mentum  chitinized;  anterior  margin  with  seven  teeth,  behind  median  tooth 
a  brush  of  hairs.  Antenna  two-segmented;  basal  segment  cylindrical;  second  segment 
button-like,  with  apex  pointed.  Mandible  three- toothed;  a  tuft  of  hairs  at  prosthecal  region. 
Spiracular  disk  surrounded  by  two  fleshy  welts  which  are  deeply  incised  medially  to  appear 
as  four  blunt  lobes.  Spiracles  difficult  to  distinguish  in  a  position  of  rest,  elliptical,  margined 
with  blackish  brown. 

Pupa. —  Length,  8  mm. 
Width,  1.75  mm. 


854  CHARLES  PAUL  ALEXANDER 

Coloration  blackish;  abdomen  dark  greenish  gray. 

Pupa  (Plate  XLI,  181)  attached  to  old  larval  skin,  which  in  turn  remains  attached  to 
rock  surface.  (Just  before  pupating,  the  larva  sticks  itself  to  the  surface  by  means  of  its 
saliva.)  Pronotal  breathing  horns  cylindrical,  a  little  swollen  at  ends.  Sheaths  of  append- 
ages dull  and  black  in  color.  Leg  sheaths  attaining  end*of  fourth  abdominal  segment. 

Dactylolabis  wodzickii  (Now.) 

'  1867    Rhicnoptila  wodzickii  Now.    Verh.  Zool.-Bot.  Ges.  Wien,  vol.  17,  p.  337-354. 

The  species  Dactylolabis  wodzickii  was  first  found  in  the  high  Alpine 
region  of  the  Hungarian  Tatras,  at  an  altitude  of  from  6000  to  8000  feet, 
in  a  region  frequented  by  the  wall  creeper  (Tichodroma).  Here  the 
adults  and  the  immature  stages  were  found  together  on  the  granite  cliffs 
where  water  dripped  continuously,  supporting  a  considerable  algal  flora. 
Associated  with  the  species  were  other  flies,  Liancalus  virens  (Scop.)  and 
Clinocera  fontinalis  Hal.,  as  well  as  Tricyphona  schineri  (KoL).  The 
degenerate  wings  of  the  insects  prevent  their  flying,  and  they  probably 
live  and  die  close  to  the  place  where  the  eggs  are  deposited.  One  fly  was 
observed  laying  her  eggs  in  the  masses  of  algae  in  this  situation.  The 
larvae  live  in  these  masses  and  pupate  in  them. 

Larva. —  Length,  12.5  mm. 
Width,  2.4  mm. 

Dorsum  of  body  dark  greenish  brown,  with  dark  longitudinal  brown  stripes  producing  a 
curious  pattern  of  straight  and  convergent  lines;  ventral  surface  almost  white,  unmarked. 

Body  depressed,  flattened,  abdominal  segments  divided  into  a  narrow  basal  and  a  much 
broader  posterior  annulus.  Head  capsule  (Plate  XLI,  185)  compact,  massive.  Mandible 
showing  but  a  single  inner  tooth.  Posterior  abdominal  annuli  with  two  long  setae  on  lateral 
margins.  Spiracular  disk  (Plate  XLT,  186)  surrounded  by  four  lobes,  t»he  ventral  pair  longer 
than  the  others;  these  lobes  margined  with  darker  chitinized  lines  and  provided  with  fringes 
of  moderately  long  hairs.  Spiracles  not  described,  and  undoubtedly  overlooked  by  the 
describer. 

Pupa.—  Length,  12.5-13.5  mm. 
Width,  2.6  mm. 

Color  brown.  Pronotal  breathing  horns  short,  cylindrical.  Wing  sheaths  ending  opposite 
middle  of  second  abdominal  segment.  Leg  sheaths  ending  beyond  middle  of  third  segment. 
Abdomen  armed  with  rows  of  spines  (Plate  XLI,  184). 

Dactylolabis  cubitalis  (O.  S.) 

1869    Limnophila  cubitalis  O.  S.    Mon.  Dipt.  N.  Amer.,  part  4,  p.  229. 
Dactylolabis  cubitalis  is  a  local  species,  often  occurring  in  great  numbers. 
In  the  gorge  of  Fall  Creek  near  the  Cornell  University  campus  this  species 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  855 

can  be  found  in  myriads  by  sweeping  the  rank  herbage  in  May.  The 
writer  has  never  located  the  immature  stages,  but  the  species  was  reared 
by  Dr.  J.  G.  Needham  at  Ithaca,  New  York,  on  May  20,  1898.  There 
is  no  record  as  to  where  these  larvae  occurred,  but  they  are  presumably 
mud-inhabiting  forms,  or  possibly  they  frequent  habitats  similar  to  those 
described  for  the  two  preceding  species. 

Pupa.' —  Length  of  cast  skin,  about  12  mm. 

Pronotal  breathing  horns  (Plate  XLI,  187)  short,  cylindrical,  the  tips  scarcely  enlarged. 
Mesonotum  behind  with  a  prominent  rounded  tubercle  on  either  side  of  median  line,  which 
is  set  with  two  spines,  a  larger  outer  spine  and  a  small  inner  spine.  Wing  sheaths  ending 
opposite  apex  of  second  abdominal  segment.  Leg  sheaths  ending  opposite  apex  of  third 
abdominal  segment,  the  tarsal  sheaths  terminating  on  a  common  level. 

Abdominal  segments  divided  into  two  rings.  Tergites  with  posterior  ring  having  a 
transverse  row  of  elongate  tubercles,  there  being  about  eight  on  segment  2,  about  six  on 
intermediate  segments  (3  to  5),  and  about  four  on  segments  6  and  7;  these  tubercles  fleshy, 
but  sparsely  armed  with  chitinized  spines.  Basal  ring  unarmed.  Pleura  with  a  strong 
tubercle  on  each  ring;  a  blunt  but  prominent  spiracle  at  extreme  base  of  posterior  ring;  this 
spiracle  very  prominent  on  segment  2  (Plate  XLI,  188),  much  exceeding  lateral  tubercles 
and  being  about  two-fifths  length  of  pronotal  breathing  horns.  Sternites  with  about  six 
tubercles  on  posterior  ring.  Male  cauda  (Plate  XLI,  189)  with  tergal  lobes  slender,  running 
out  into  acute  chitinized  points,  which  are  directed  caudad,  slightly  divergent;  dorsum  of 
segment  8  with  posterior  lobes  powerful,  about  equal  in  size  to,  or  a  little  larger  than,  tergal 
lobes  of  cauda  just  described;  anterior  lobes  small,  directed  caudad  and  laterad;  two  large 
lateral  lobes  and  two  smaller  ventral  lobes.  Female  cauda  with  tergal  valves  moderately 
elongate,  stout,  broad  at  base,  narrowed  toward  tip,  where  they  run  out  into  long,. chitinized 
points  directed  caudad  and  slightly  dorsad.  ^ 

Neanotype  —  Cast  pupal  skin,  Ithaca,  New  York,  May  20,~1898.    - 
Paratype. —  Pupae,  cast  skins,  with  type. 

Dactylolabis  montana  (0.  S.) 

1859  Limnophila  montana  0.  S.  Proc.  Acad.  Nat.  Sci.  Phila.,  p."240r^  ^ 
The  species  Dactylolabis  montana  is  common  and  widely  distributed 
thruout  the  northern  United  States.  It  is  a  characteristic  inhabitant  of 
rocky  cliffs,  where  it  rests  in  crevices  on  the  almost  vertical  walls.  The 
immature  stages  have  long  remained  unknown,  but  during  the  spring 
of  1920  W.  L.  McAtee  found  a  pupal  skin  and  the  teneral  adult  near 
Washington,  D.  C.  Mr.  McAtee  writes  that  the  pupa  was  found  in  the 
moss  that  covers  the  almost  vertical  north  fac3  of  a  cliff  on  Plummers 
Island;  this  moss  is  mostly  shaded,  and  grows  on  a  thin  layer  of  black  soil 
which,  at  that  time  of  the  year  at  least,  is  saturated  with  water.  The 


856  CHARLES  PAUL  ALEXANDER 

pupal  skin  was  not  in  condition  for  study,  but  now  that  the  larval  habitat 
is  known  it  is  hoped  that  more  material  of  the  immature  stages  will 
become  available. 

Subtribe  Limnophilaria 

The  division  Limnophilaria  comprises  a  large  and  heterogeneous  group 
of  species  which  in  their  larval  and  pupal  characters  grade  rather  insensibly 
into  the  next  subtribe,  the  Hexatomaria,  altho  the  adults  of  the  two 
subtribes  are  very  distinct. 

The  larvae  of  the  various  species  in  the  group  Limnophilae  are  rather 
similar  to  one  another.  The  head  capsule  is  long  and  narrow,  with  the 
lateral  plates  very  slender.  The  labrum  is  ample,  with  several  sensory 
papillae  and  setae.  The  antennae  (which  are  two-segmented  in  the 
subgenera  Phylidorea  and  Lasiomastix,  and  probably  also  in  Dicrano- 
phragma  and  others)  bear  at  their  tips,  in  addition  to  the  usual  elongate 
setae,  an  oval  or  elongate-oval  papilla  which  is  delicately  sculptured. 
The  mental  region  is  feebly  chitinized,  and  consists  of  two  short  longitudinal 
bars  articulated  at  their  cephalic  ends  with  a  transverse  bar  which  functions 
as  the  mental  plate;  this  bar,  or  plate,  is  usually  delicately  grooved  with 
parallel  striae,  a  type  of  articulation  that  allows  for  great  distention  of 
the  gular  region  in  feeding.  The  mandibles  are  powerful  chitinized  hooks 
bearing  two  or  more  acute  or  flattened  teeth  at  about  midlength.  The 
outer  lobes  of  the  maxillae  project  far  cephalad  as  pale  flattened  blades. 

The  spiracular  disk,  in  the  primitive  condition,  is  squarely  truncated 
and  is  surrounded  b}^  five  subequal  lobes  which  are  fringed  with  rather 
short  hairs.  The  dorso-median  lobe  is  lost  in  most  species,  but  the  two 
pairs  of  lobes  are  more  or  less  preserved  in  the  other  species  known  to  the 
writer,  altho  the  lateral  pair  is  sometimes  reduced  to  a  mere  fringe  or 
tuft  of  hairs.  The  terminal  fringes  of  the  ventral  lobes  are  often  greatly 
elongated,  and  fanlike.  The  anal  gills  are  four  in  number,  retractile, 
and  rarely  conspicuous.  As  is  usual  in  many  mud-  or  sand-inhabiting 
species  of  Tipulidae,  the  larvae  are  capable  of  greatly  distending  the 
subterminal  abdominal  segment.  This  segment  is  often  provided  with 
numerous  transverse  rows  of  fine  points,  and  its  inflation  assists  in  the 
larva's  progression  thru  the  soil. 

The  larvae  of  the  group  are  carnivorous,  and  almost  without  exception 
are  exceedingly  agile  and  snakelike  in  their  motions. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  857 

The  pupae  of  the  group  Limnophilae  have  the  pronotal  breathing  horns 
usually  small,  or  at  least  not  greatly  elongated.  The  abdomen  is  often 
greatly  depressed,  with  the  lateral  margins  flattened  and  carinate  and 
the  segments  incised.  In  the  species  of  the  subgenus  Phylidorea  the 
segments  of  the  abdomen  are  armed  with  rows  of  long,  slender  spines. 

The  group  Ulomorphae  seems  to  be  closely  related  to  the  other  members 
of  the  division,  but  the  immature  stages  present  some  curious  conditions 
which  it  is  difficult  to  correlate  with  the  same  structures  in  the  group 
Limnophilae.  The  writer  has  removed  a  group  of  species  from  the  old 
genus  Limnophila,  and  has  placed  these  in  the  genus  Pilaria  Sintenis. 

In  the  larvae  of  the  group  Ulomorphae,  the  epipharynx  and  the  maxillae 
are  densely  hairy.  The  maxillary  lobes  are  slender.  The  mandibles 
are  hinged  at  about  midlength,  the  basal  segment  being  concave  on  its 
inner  face  to  receive  the  mandibular  blade  when  in  a  position  of  rest. 
The  mandibular  blade  has  one,  or  sometimes  two,  elongate  acute  teeth 
at  the  base.  The  mental  region  is  not  chitinized.  The  spiracular  disk 
is  surrounded  by  four  lobes,  of  which  the  lateral  pair  are  often  very 
reduced;  the  inner  faces  of  the  lateral  lobes  are  capable  of  close  approxi- 
mation, so  that  when  they  are  closely  applied  the  spiracles  are  contiguous. 
The  pupae  have  the  pronotal  breathing  horns  very  elongate,  with  their 
tips  split  into  two  flattened  divergent  lobes,  as  in  Pseudolimnophila  and 
a  few  tipuline  forms. 

The  following  keys  separate  the  genera  and  the  subgenera  of  the  subtribe 

Limnophilaria: 

Larvae 

1.  Mental  region  not  chitinized;  maxillae  and  epipharynx  fringed  with  conspicuous,  long, 

golden-yellow  hairs;  mandibles  hinged;  head  capsule  with  dorsal  plate  spatulate  at 

its  tip.     (Group  Ulomorphae,  p.  869J 2 

Mental  region  a  narrow,  transverse^  chitinized  bar,  finely  striate;  maxillae  and 
epipharynx  without  conspicuous  hairs;  mandibles  not  hinged;  head  capsule  not  as 
described  above.  (Group  Limnophilae,  p.  858) 3 

2.  Length  under  12  mm. ;  basal  tooth  of  mandibular  blade  nearly  half  the  length  of  blade. 

Ulomorpha  0.  S.  (p.  869) 

Length  over  14  mm. ;  basal  tooth  of  mandibular  blade  about  one-third  or  less  the  length 
of  blade Pilaria  Suit.  (p.  872) 

3.  Spiracular  disk  almost  squarely  truncated,  surrounded  by  five  subequal  lobes  producing 

an  eriopterine  appearance Dicranophragma  O.  S.  (p.  861) 

Spiracular  disk  obliquely  truncated,  surrounded  by  two  pairs  of  lobes  of  which  the  lateral 

pah*  are  the  shorter t -. 4 

L  Mandibles  with  two  or  three  acute  teeth  at  about  midlength;  epipharynx  with  a  circular 

area  bearing  two  biarticulate  papillae Phylidorea  Bigot  (p.  866) 

Mandibles  with  three  or  four  flattened  and  truncated  teeth  along  blade;  epipharynx 

without  papillae  as  above Lasiomastix  O.  S.  (p.  863) 


858  CHARLES  PAUL  ALEXANDER 

Pupae 

1.  Pronotal  breathing  horns  very  long,   cylindrical,   the  tips  split  into  divergent  flaps. 

(Group  Ulomorphae,  p.  869) 2 

Pronotal  breathing  horns  not  elongate-cylindrical.     (Group  Limnophilae,  p.  858) 3 

2.  Abdominal  segments  with  tubercles  or  spines  only  near  posterior  margins  of  segments. 

Ulomarpha  O.  S.  (p.  869) 
Abdominal  segments  with  three  or  four  pairs  of  blunt,  naked  tubercles. 

Pilaria  Sint.  (p.  872) 

3.  Abdominal  segments  depressed,   lateral   margins   flattened,   carinate,    segments  deeply 

incised 4 

Abdominal  segments  not  depressed  nor  incised;  pronotal  breathing  horns  bicolored. 

Dicranophragma  O.  S.  (p.  861) 

4.  Abdominal  segments  with  rows  of  acute  slender  spines;  pronotal  breathing  horns  very 

short,  semicircular  in  outline,  bluntly  rounded  at  their  tips.  .  Phylidorea  Bigot  (p.  866) 

Abdominal  segments  without  such  slender  spines;  pronotal  breathing  horns  short,  broad, 

slightly  compressed,  tips  a  little  pointed Lasiomastix  O.  S.  (p.  863) 

Group  Limnophilae 

Genus  Limnophila  Macquart  (Gr.  swamp  +  friend) 

1834    Limnophila  Macq.     Suit,  a  Buff.,  vol.  1,  Hist.  Nat.  Ins.,  Dipt.,  p.  95. 
1854     Phylidorea  Bigot.     Ann.  Soc.  Ent.  France,  p.  456. 
1861    Limnomya  Rond.     Dipt.  Ital.  Prodr.,  vol.  4,  p.  11. 

Larva. —  Form  slender.  Spiracular  disk  surrounded  by  five,  or  more  commonly  four, 
lobes,  the  ventral  pair  the  longer  and  fringed  with  long,  delicate  hairs.  Anal  gills  four,  not 
conspicuous.  Head  capsule  flattened,  very  dissected,  with  plates  narrow.  Mandible  not 
hinged,  curved,  chitinized,  on  cutting  edge  with  a  few  acute  or  flattened  teeth.  Maxilla 
with  outer  lobe  produced  into  a  flattened  projecting  blade.  Antenna  two-segmented,  at 
its  tip  with  a  short  or  elongate-oval  papilla  which  is  delicately  sculptured.  Mentum  a 
transverse  chitinized  crossbar  which  is  finely  grooved. 

Pupa.- —  Cephalic  crest  setiferous.  Pronotal  breathing  horns  usually  small,  flattened, 
tips  not  split.  Mesonotum  convex  or  flattened  above.  Abdominal  segments  in  some  cases 
armed  with  tubercles  or  spines. 

The  genus  Limnophila  comprises  a  very  extensive  group  of  crane-flies 
(more  than  one  hundred  and  fifty  species),  which  are  found  in  most  parts 
of  the  world  but  are  apparently  more  numerous  in  the  temperate  regions. 
The  adult  flies  seem  to  be  closely  related,  but  the  immature  stages  are  so 
varied  in  structure  as  to  make  it  appear  that  the  group  must  be  a  hetero- 
geneous one  and  the  similarity  of  the  adults  a  result  of  convergent  evolution. 

The  adult  flies  may  be  found  resting  on  rank  vegetation.  Several  of 
the  species  (Limnophila  ultima,  for  example)  swarm  in  small  groups  of 
from  fifty  to  sixty  individuals,  copulation  taking  place  in  the  air,  as  is 
discussed  more  fully  on  page  711. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  859 

The  immature  stages  of  several  species  have  been  made  known,  but 
there  are  still  very  considerable  gaps  in  the  present  knowledge.  The 
larvae  are  among  the  most  carnivorous  of  all  crane-fly  larvae.  The 
feeble  chitinization  of  the  montal  region  allows  for  great  distention,  and 
oftentimes  the  prey  of  these  larvae  consists  of  forms  that  are  nearly  as 
large  as  the  captors  themselves. 

The  data  regarding  extra-American  species  may  be  summarized  as 
follows : 

Limnophila  bryobia  Mik  (1881:205-206)  was  reared  from  moss,  taken  in 
the  Auckland  Islands,  near  New  Zealand,  in  the  summer  (December  to 
February)  of  1874-75.  The  moss  was  dried  and  then  placed  in  a  tem- 
perature of  from  12°  to  15°  Reaumur.  In  March  of  1879  an  adult  fly 
appeared.  This  long  interval  of  more  than  four  years  was  supposed  by 
Mik  to  have  been  passed  by  the  insect  as  a  larva,  the  dry  moss  furnishing 
the  food.  Possibly  the  eggs  were  carried  over  for  a  long  period  before 
hatching. 

Limnophila  sinistra  Hut  ton,  of  New  Zealand,  has  been  discussed  in  some 
detail  by  Hudson  (1920:33-34),  who  states  that  this  species  is  Common  in 
most  densa  forests  thruout  the  country.  The  larva  inhabits  fallen  tree- 
trunks  in  an  advanced  state  of  decay,  forming  burrows  between  the  soft, 
decayed  part  and  the  harder  part  of  the  wood.  It  is  about  25  millimeters 
in  length,  cylindrical,  tapering  toward  the  head,  which  is  very  small,  and 
furnished  with  two  minute  jaws  and  a  pair  of  very  short  antennae.  There 
are  eleven  visible  body  segments.  The  extremity  of  the  last  segment  is 
truncate  and  deeply  excavated,  the  concavity  being  protected  by  five  con- 
verging spines,  which  can  be  spread  out  or  drawn  inward  at  the  will  of 
the  insect.  Pedal  warts  occur  on  the  undersides  of  all  the  segments  except 
the  three  immediately  following  the  head  and  the  terminal  segment.  The 
pupa  is  about  12  millimeters  in  length,  and  rather  stout.  The  breathing 
horns  are  about  one-third  the  length  of  the  wing  sheaths,  and  are  mod- 
erately stout  and  strongly  recurved.  There  are  two  dorsal  rows  of  hooks 
on  each  exposed  abdominal  segment  and  one  ventral  row  near  the  termi- 
nal extremity.  The  valves  of  the  ovipositor  are  strongly  recurved  and 
very  stout.  The  head  and  the  thorax  are  dark  brownish  black,  highly 
polished;  the  abdomen  is  grayish  ocherous,  darker  in  the  middle;  the  ovi- 
positor arid  the  terminal  segments  of  the  posterior  tarsi  are  reddish. 
The  pupa  rests  in  a  burrow  made  by  the  larva  near  the  surface  of  the  log. 


860  CHARLES  PAUL  ALEXANDER 

1  .  i 

Edwards  has  decided  that  this  insect  and  a  few  related  species  should  be 
placed  in  a  new  genus.  The  affinities  of  the  fly,  to  judge  from  the  details 
supplied  by  Hudson,  are  with  the  Epiphragmaria  rather  than  with  the 
Limnophilaria,  but  until  more  details  are  forthcoming  it  cannot  be  defi- 
nitely referred  to  that  division. 

Limnophila  (Phylidorea)  ferruginea  (Meig.),  of  Europe,  is  closely  allied 
to  L.  adusta  (page  867),  and  is  of  especial  interest  as  being  the  type  of 
the  subgenus  Phylidorea.  It  was  reared  by  De  Meijere  (1916:204-206) 
from  larvae  found  living  between  saturated  decaying  leaves  along  the 
banks  of  watercourses.  Further  mention  of  the  species  is  made  under 
the  discussion  of  the  subgenus  Phylidorea  (page  866). 

Limnophila  hyalipennis  (Zett.),  of  Europe,  was  found  by  Beling  (1886: 
198-199)  in  piny  woods,  where  the  larvae  live  in  old  ant  hills  or  in  piles 
of  earth  thrown  up  by  wagon  wheels.  The  apparent  lack  of  pupal  breathing 
horns  in  this  species  is  discussed  on  page  755.  The  European  species 
L.  lineola  (Meig.)  (Beling,  1879:54  and  1886:199-200),  L.  nemoralis 
(Meig.)  (Beling,  1886:200-201),  and  L.  ochracea  (Meig.)  (Beling,  1886- 
202)  are  all  found  in  damp  earth  along  the  margins  of  woodland  streams 
or  in  damp  spots  in  the  woods.  Larvae  of  L.  dispar  (Meig.)  were  found 
by  Perris  (1849:331)  living  in  the  hard,  withered  stalks  of  an  umbellifer, 
Angelica  sylvestris  Linn.,  where  they  hollowed  out  long  passages  in  the 
pith.  (This  reference  is  cited  by  Mik,  1881 : 204,  and  by  Osten  Sacken, 
1869:201-202.)  Limnophila  pallida  Bel.  was  reared  by  Beling  (1873 a: 
557)  from  larvae  living  in  a  decaying  ash  trunk. 

The  reference  to  Limnophila  platyptera  Macq.  given  by  Heeger  (1854) 
is  considered  by  Osten  Sacken  (1869:4,  202)  to  refer  to  a  mycetophilid, 
probably  Bolitophila. 

The  American  species  that  are  now  known  are  considered  in  the 
following  pages.  In  addition,  Limnophila  unica  0.  S.  has  been  bred  from 
larvae  found  in  decaying  wood. 

It  is  very  probable  that  some  of  the  species  listed  above,  as  well  as 
many  of  the  limnophiline  forms  that  are  still  unknown  as  regards  their 
immature  stages,  will  be  found  to  belong  to  some  one  or  other  of  the 
remaining  divisions  of  the  tribe,  rather  than  to  the  Limnophilaria  as  here 
restricted.  This  group  requires  more  careful  study  than  does  any  other 
division  of  crane-flies. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  861 

(Subgenus  Dicranophragma  Osten  Sacken) 

1859     Dicranophragma  0.  S.    Proc.  Acad.  Nat.  Sci.  Phila.,  p.  240. 

Larva. —  Form  short  and  stout.  Spiracular  disk  surrounded  by  five  short  lobes,  the 
ventral  pair  the  longest,  the  dorsal  lobe  very  blunt;  disk  surrounded  by  a  fringe  of  delicate 
hairs  which  are  longest  at  tips  of  lobes.  Head  capsule  of  the  hexatomine  type.  Mandible 
powerful,  prolonged  into  a  strong  hook  with  a  large  acute  tooth  just  beyond  midlength. 
Maxilla  very  long  and  slender.  Mentum  chitinized,  consisting  of  a  narrow  transverse  bar 
which  is  ribbed  with  fine  parallel  grooves. 

Pupa. —  Cephalic  crest  prominent,  each  half  with  three  setae.  Pronotal  breathing  horns 
short,  cylindrical,  or  slightly  narrowed  to  a  blunt  tip.  Mesonotum  convex,  unarmed. 
Wing  sheaths  short,  broad,  ending  before  tip  of  second  abdominal  segment.  Leg  sheaths 
moderate  in  length,  ending  just  beyond  base  of  fourth  abdominal  segment,  hind  tarsi  a  little 
longer  than  the  others.  Abdominal  segments  divided  into  two  rings;  posterior  ring  with 
a  subterminal  transverse  armature,  on  dorsum  consisting  of  numerous  setiferous  tubercles, 
on  pleura  consisting  of  two  or  three  groups  of  setiferous  tubercles;  basal  ring  with  about 
eight  slender  tubercles  on  dorsal  surface.  Spiracles  present,  but  evidently  nonfunctional. 
Dorsum  of  segment  8  with  four  slender  tubercles. 

Dicranophragma  is  a  well-defined  group  of  the  genus  Limnophila,  with 
about  six  known  species  whose  center  of  distribution  seems  to  be  in  the 
Oriental  region.  The  type  of  the  subgenus,  Limnophila  (Dicranophragma) 
fuscovaria,  discussed  in  detail  below,  is  the  only  described  New  World 
form.  The  immature  stages  are  spent  in  rich  organic  mud  in  cool, 
shaded  woods. 

Limnophila  (Dicranophragma)  fuscovaria  0.  S. 

1859    Limnophila  (Dicranophragma)  fuscovaria  0.  S.    Proc.    Acad.    Nat.    Sci.    Phila., 
p.  240. 

The  adult  flies  of  Limnophila  fuscovaria  may  be  easily  distinguished  by 
their  broad,  heavily  spotted  wings  and  the  strong  supernumerary  cross  vein 
in  cell  R%.  They  may  be  swept  from  rank  vegetation  in  cool,  shaded 
woods.  The  larva  has  the  body  stouter  and  less  elongate  than  is  usual 
in  this  group  of  species,  but  it  possesses  the  same  snakelike  movements 
as  its  near  relatives.  The  pupal  duration  is  not  longer  than  eight  days 
(June  8  to  16,  in  1917). 

Larva. —  Length,  6.8-7.2  mm. 
Djameter,  0.7-0.8  mm 

Coloration  light  golden-yellow;  maxillary  lobes  yellow;  spiracular  disk  lined  with  dark  brown. 
Body  rather  short  and  stout,  not  of  the  exceedingly  elongate  type  of  the  Eriopterini,  gradu- 
ally narrowed  to  anterior  end  (Plate  XLII,  190) ;  prothoracic  segment  truncated  anteriorly; 


862  CHARLES  PAUL  ALEXANDER 

when  head  is  retracted,  tips  of  maxillae  projecting  from  prothoracic  orifice.  Seventh 
abdominal  segment  with  a  transverse  fringe  of  stiff  hairs  on  the  sides,  these  directed  laterad 
and  slightly  caudad.  Body  noticeably  constricted  just  before  last  segment.  Spiracular 
disk  (Plate  XLII,  195)  surrounded  by  five  lobes,  and  in  superficial  appearance  decidedly 
eriopterine  rather  than  hexatomine;  ventral  lobes  the  longest  but  still  much  shorter  than 
is  usual  in  this  tribe,  slightly  divergent,  inner  face  with  a  double,  V-shaped,  dark  brown 
mark,  inner  arm  of  V  narrow,  outer  arm  more  expanded,  especially  at  proximal  ends,  one 
large  inner  and  two  smaller  outer  sensory  bristles  at  apex  of  V;  lateral  lobes  narrow,  with  a 
V-shaped  mark  similar  to  that  of  ventral  lobes  but  narrower  and  darker;  dorsal  lobe  broad, 
the  inner  face  indistinctly  lined  with  parallel,  dusky,  longitudinal  stripes;  all  five  lobes 
surrounded  by  a  fringe  of  long,  delicate  hairs  lying  just  outside  brown  marks,  these  hairs 
longest  at  tips  of  lobes  and  continuous  around  disk.  Spiracles  circular,  rather  widely  sepa- 
rated; ring  broad,  light  yellow,  narrowly  margined  outwardly  with  black.  Anal  gills  four, 
short  and  stout. 

Head  capsule  (Plate  XLII,  191)  very  dissected,  consisting  of  six  elongate  slender  plates, 
three  on  either  side,  articulating  at  a  point  laterad  of  mental  plate.  Labrum  (Plate  XLII, 
192)  large;  anterior  margin  indistinctly  trilobed,  the  median  lobe  hairy;  on  the  ventral,  or 
epipharyngeal,  surface,  on  either  side,  a  slender  cylindrical  papilla,  subhyaline,  directed 
cephalad;  sides  of  epipharynx  with  a  tuft  of  long  setae.  Mentum  chitinized,  consisting  of 
two  lateral  rods  and  an  anterior  transverse  bar  which  is  finely  ribbed  with  parallel  lines. 
Immediately  dorsad  of  mentum  two  roughened  papillae  (prementum)  directed  cephalad; 
esophagus  retrorsely  roughened.  Antenna  (Plate  XLII,  193)  stout,  in  caustic-potash  prepa- 
rations almost  hyaline,  tipped  with  an  elongate  cylindrical  lobe  distad  of  which  is  a  small 
setiferous  tubercle  with  two  long,  delicate  setae  exceeding  antennae  in  length.  Mandible 
(Plate  XLII,  194)  powerful,  very  deep  at  base,  produced  into  an  acute,  strongly  curved  tip; 
just  beyond  midlength  a  very  large,  acute  tooth  which  is  about  one-third  length  of  apical 
point,  in  its  angle  a  small,  hyaline,  flattened,  leafiike  blade  with  a  truncated  apex;  basad 
of  large  tooth  two  or  three  gradually  smaller  teeth.  Maxilla  with  outer  lobes  stout,  broad 
at  base,  gradually  narrowed  to  tip,  which  is  cut  off  by  a  constriction  into  a  semi-oval  struc- 
ture; palpi  on  inner  dorsal  face  of  maxillary  lobe  at  about  midlength,  short,  semi-globular, 
with  abundant  sensory  papillae;  when  head  is  completely  retracted,  the  long,  pale,  slightly 
divergent  tips  of  maxillary  lobes  projecting  from  orifice. 

Pupa. —  Length  of  cast  skin,  about  6  mm. 

General  coloration  dark  brown;  pronotal  breathing  horns  almost  black,  with  apical  fourth 
conspicuously  light  yellow;  abdominal  incisures  pale. 

Cephalic  crest  large  and  prominent,  with  three  long  setae  on  outer  face.  Labrum  blunt 
at  apex.  Labial  lobes  with  a  blunt  point  on  either  side,  these  directed  proximad  and  caudad. 
Maxillary  palpi  broad  at  base,  suddenly  narrowed  to  the  long,  cylindrical  tip,  which  is 
darkened.  Antennae  rather  short.  Pronotal  breathing  horns  (Plate  XLIII,  196)  stout, 
short,  almost  straight,  broad  basally,  tapering  gradually  to  the  blunt  tips.  Wing  sheaths 
very  short  and  broad,  ending  before  tip  of  second  abdominal  segment.  Leg  sheaths  short, 
extending  to  just  beyond  base  of  fourth  abdominal  segment;  all  the  tarsi  ending  about  on  a 
level,  or  outer  sheaths  a  little  the  longer.j 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  863^ 

Abdominal  segments  (Plate  XLIII,  198)  each  divided  into  two  rings  by  a  false  constriction, 
basal  ring  about  two- thirds  length  of  posterior  ring.  Basal  ring  on  dorsum  with  a  transverse 
row  of  about  eight  slender,  blunt  tubercles  before  caudal  margin;  on  pleura  a  similar  but 
larger  tubercle  opposite  basal  ring,  this  tipped  with  a  long,  delicate  seta.  Sternum  unarmed. 
Posterior  ring  on  dorsum  with  a  subterminal  row  of  blunt  tubercles,  simple  or  bifid,  often 
approximated  or  crowded,  bearing  one  or  two  long  setae;  near  base  of  tergum,  opposite 
spiracle,  a  setiferous  tubercle;  on  pleura,  opposite  base  of  posterior  ring  and  nearer  dorsum 
than  sternum,  a  vestigial  spiracle  with  a  double  setiferous  tubercle  ventrad  of  it;  near  caudal 
margin  of  pleura  two  or  three  compound  multisetose  tubercles  (Plate  XLIII,  197)  or  aggre- 
gations of  simple  tubercles,  which  are  largest  on  seventh  segment.  On  sternum,  two  slender 
tubercles  on  either  side,  near  base  of  posterior  ring.  Female  cauda  (Plate  XLIII,  201) 
elongate,  with  the  tergal  valves  a  little  longer  than  the  sternal  valves;  each  of  tergal  valves 
with  a  small  tubercle  at  base.  Male  cauda  (Plate  XLIII,  199  and  200)  with  the  ventral 
lobes  blunt,  the  dorsal  lobes  a  little  longer  and  ending  in  an  acute  point;  a  tiny  tubercle 
on  either  side  at  base  of  dorsal  lobes.  Dorsum  of  segment  8  with  a  trapezoid  of  four  con- 
spicuous lobes  ending  in  truncated  tubercles;  anterior  pair  of  lobes  smaller  and  a  little  more 
widely  separated  than  posterior  pair. 

Nepionotype. —  Ithaca,  New  York. 

Neanotype  —  Ithaca,  New  York,  May  31,  1917.     No.  57-1917. 

Paratypes.—  Pupal  skins,  June  11,  1917  (No.  99-1917),  June  16,  1917  (No.  132-1917). 
Abundant  larvae,  with  nepionotype. 

(Subgenus  Lasiomastix  Osten  Sacken) 

1859    Lasiomastix  0.  S.    Proc.  Acad.  Nat.  Sci.  Phila.,  p.  233. 

Larva. —  Body  slender,  with  pencils  of  stiff  setae  on  lateral  margins  of  thorax.  Spiracular 
disk  with  four  lobes,  the  ventral  pair  a  little  the  longer  and  bearing  a  fringe  of  long,  delicate 
hairs,  these  about  twice  length  of  lobes.  Anal  gills  short  and  stout.  Head  capsule  of 
hexatomine  type.  Mandible  powerful,  cutting  edge  with  a  single  row  of  three  or  four 
flattened,  bladelike  teeth.  Maxilla  very  long  and  slender.  Antenna  elongate,  at  tip  with 
a  small  elongate-oval  papilla  whose  surface  is  delicately  sculptured.  Mentum  chitinized, 
a  strong  transverse  bar  which  is  ribbed  with  fine  parallel  grooves. 

Pupa.—  Cephalic  crest  very  small,  inconspicuous,  bilobed.  Pronotal  breathing  horns 
short,  broad,  somewhat  compressed,  a  little  pointed  at  apex.  Mesonotum  unarmed. 
Abdominal  segments  depressed,  armed  with  numerous  tubercles  or  small  spines,  these  being 
most  abundant  on  posterior  annuli  of  segments. 

The  subgenus  Lasiomastix  includes  but  six  known  species,  three  of 
which  are  from  eastern  North  America.  Limnophila  (Lasiomastix) 
macrocera  is  common  and  widely  distributed  thruout  the  Eastern  States. 
L.  (L.)  tenuicornis  O.  S.  and  L.  (L.)  subtenuicornis  (Alex.)  are  found  only 
in  the  Northeastern  States  and  are  more  local  in  their  distribution.  The 
immature  stages  of  L.  (L.)  macrocera  are  spent  in  rich  organic  mud.  They 
are  very  similar  to  those  of  the  subgenus  Phylidorea, 


864  CHARLES  PAUL  ALEXANDER 

Limnophila  (Lasiomastix)  macrocera  (Say) 

1823    Limnobia  macrocera  Say.    Journ.  Acad.  Nat.  Sci  Phila.,  vol.  3,  p.  20. 

Limnophila  macrocera  is  a  common  swamp-inhabiting  crane-fly,  the 
larvae  living  in  organic  mud.  At  Orono,  Maine,  the  writer  found  larvae 
of  this  species  associated  with  larvae  of  Bittacomorpha  davipes,  Pilaria 
tenuipes,  P.  recondita,  Erioptera  chlorophylla,  and  other  crane-flies,  as 
well  as  with  leeches,  snails,  worms  of  many  kinds,  and  other  forms  of 
life.  The  larva  is  similar  to  that  of  other  related  species.  When  placed 
in  water  it  is  very  active  and  has  the  habit  of  darting  the  anterior  quarter 
of  its  body  from  one  side  to  the  other,  suggesting  the  striking  of  a  reptile. 
The  pupal  duration  is  not  more  than  eight  days  (June  24  to  July  2,  June 
28  to  July  6). 

Larva. —  Length,  14.5-15  mm. 
Diameter,  1.4  mm. 

Coloration,  pale  yellowish  white. 

Body  terete,  narrowed  toward  both  ends  but  more  noticeably  toward  anterior  end. 
Integument  covered  with  a  dense  appressed  pubescence.  Chaetotaxy  as  follows:  two  stout 
setae  on  both  dorsal  and  ventral  margins  of  prothoracic  orifice;  thorax  with  pencils  of  stiff 
setae  near  anterior  margins  of  segments,  two  large  lateral  tufts  and  a  smaller  ventral  pair; 
abdominal  segments  with  a  pencil  of  similar  bristles  on  ventro-lateral  margins,  one  on 
anterior  half,  the  other  midlength,  of  each  ring;  a  seta  at  posterior  lateral  angles  of  sternites; 
four  setae  in  a  transverse  row  near  posterior  margin  of  tergites,  the  lateral  pair  the  larger; 
lateral  margins  of  cauda  near  base  of  lateral  lobes  and  above  gills  with  pencils  of  blackish 
setae;  basal  part  of  segments  on  both  sternum  and  tergum  with  transverse  parallel  rows  of 
fine  scabrous  points,  this  area  very  narrow  on  second  abdominal  segment,  consisting  only 
of  three  or  four  rows,  the  areas  gradually  becoming  broader  and  the  rows  more  numerous, 
there  being  on  sixth  and  seventh  segments  about  28  to  30  rows  which  occupy  nearly  the 
basal  third  of  segments. 

Spiracular  disk  ( Plate  XLIV,  208)  surrounded  by  four  lobes;  ventral  pair  the  longest,  lateral 
pair  a  little  shorter;  ventral  lobes  near  tips  with  a  brush  of  delicate,  exceedingly  long  hairs 
which  curl  into  loops  at  tips,  the  longest  of  these  hairs  about  twice  length  of  lobes  bearing 
them;  this  fringe  of  hairs  continuous  around  disk,  longest  at  ends  of  lobes,  gradually  shortened 
toward  their  bases  but  not  interrupted;  lateral  lobes  with  terminal  hairs  a  little  shorter  but 
still  longer  than  lobes  themselves;  inner  faces  of  lobes  delicately  margined  with  dark  brown, 
those  of  ventral  lobes  having  the  outer  lateral  margins  the  broadest.  Spiracles  rather  small, 
located  at  base  of  lateral  lobes.  Anal  gills  four,  very  stout  and  plump. 

Head  capsule  with  a  framework  of  long,  slender  plates,  as  is  usual  in  this  group.  Labrum 
(Plate  XLIV,  205)  broadly  transverse;  anterior  margin  irregularly  rounded;  anterior  median 
area  truncated,  with  the  lateral  angles  slightly  projecting,  cut  off  squarely,  each  with  a  small, 
hyaline,  sensory  papilla;  laterad  and  caudad  of  each,  two  sensory  papillae,  the  innermost 
elongate-cylindrical,  more  than  twice  length  of  short,  oval,  outer  one.  Epipharyngeal 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  865 

region  roughened  by  tiny  groups  of  parallel  ridges.  Mental  region  consisting  of  a  heavily 
chitinized  transverse  bar  which  is  delicately  grooved,  enlarged  at  ends,  and  articulating  with 
a  small  ventral  bar.  Antenna  (Plate  XLIV,  206)  long  and  slender;  basal  segment  short  and 
broad;  second  segment  elongate-cylindrical,  bearing  at  its  tip  a  small,  hyaline,  sensory  papilla 
which  appears  delicately  crosshatched  by  fine  impressed  lines,  this  papilla  elongate-oval 
with  the  apex  broadly  rounded;  a  long  hair  near  base  of  this  papilla  and  about  twice  its  length. 
Mandible  (Plate  XLIV,  207)  long  and  powerful,  the  tip  produced  into  a  slender  point,  the 
cutting  edge  at  about  midlength  with  a  single  row  of  three  or  four  flattened,  bladelike  teeth 
which  are  truncated  at  their  tips,  these  teeth  varying  considerably  in  their  shape  and  relative 
proportions.  Maxilla  with  outer  lobe  elongate,  the  base  strongly  chitinized,  this  chitinized 
part  continuing  up  margin  of  lobe  almost  to  tip,  the  apical  part  nearly  hyaline;  at  about 
midlength,  the  small  palpus  and  a  long  seta.  Esophagus  enlarged,  the  walls  thickened  and 
roughened. 

(A  slide  of  a  larva  that  was  supposed  to  belong  to  this  species  differs  considerably  from 
the  material  described  above.  The  antenna  is  much  shorter,  with  the  apical  tubercle  much 
larger  and  elongate-cylindrical;  the  mandible  has  the  teeth  fewer  and  more  acute,  quite 
as  shown  by  De  Meijere  for  Limnophila  [Phylidorea]  ferruginea.  It  is  probable  that  this 
species  likewise  is  a  Phylidorea.) 

Pupa. —  Length,  14-16  mm. 

Width,  d.-s.,  1.8-2  mm. 
Depth,  d.-v.,  1.9-2.2  mm. 

Breathing  horns  pale  whitish  brown,  a  little  darker  at  base;  remainder  of  body  dark 
brownish  black,  abdomen  a  little  lighter-colored;  incisures  of  segments,  and  pleural  line,  pale. 

Cephalic  crest  very  small,  inconspicuous,  bilobed,  each  principal  lobe  subdivided  into 
two  smaller  lobes  terminating  in  a  small  seta;  another  seta  on  anterior  face.  Labrum 
rather  blunt  and  truncated  at  apex.  Labial  lobes  terminating  in  sharp  points  projecting 
proximad.  Maxillary  palpi  short  and  stout,  tapering  to  blunt  tips  (Plate  XLV,  210). 
Antenna  of  male  elongate,  reaching  to  beyond  midlength  of  wing  sheath;  antenna  of  female 
short,  ending  at  about  one-fourth  length  of  wing  sheath.  Segments  of  antenna  cylindrical, 
unarmed. 

Pronotal  breathing  horns  (Plate  XLV,  209)  short,  broad,  somewhat  compressed,  a  little 
pointed  at  tip.  Thorax  broad  and  deep,  the  pronotum  flattened,  carinate  medially. 
Mesonotum  high,  convex,  transversely  wrinkled.  Wing  sheaths  ending  some  distance  before 
tip  of  second  abdominal  segment.  Leg  sheaths  short,  ending  before  tip  of  third  abdominal 
segment,  the  hind  legs  a  little  longer  than  the  other  legs,  which  end  about  on  a  common 
level. 

Abdomen  depressed,  lateral  margin  carinate.  Abdominal  segments  distinct,  incised,  each 
segment  divided  into  two  a'nnuli,  basal  one  about  half  length  of  posterior  one.  Basal  ring 
on  dorsum  with  a  transverse  single,  or  somewhat  double,  row  of  small  subacute  spines,  which 
are  more  numerous  and  larger  on  mid-dorsal  area.  Pleura  unarmed.  Sternum  with  an 
oblique  flattened  lobe  or  wing  on  each  side,  directed  caudad  and  proximad;  between  these 
lobes  a  more  or  less  broken  transverse  row  of  from  six  to  twelve  slender  tubercles.  Posterior 
ring  on  dorsum  with  numerous  small  scattered  tubercles.  Pleura  with  a  small  circular 
spiracle  near  base  and  close  to  dorsal  margin;  a  slender  seta  near  spiracle  and  ventrad  of  it; 


866  CHAKLES  PAUL  ALEXANDER 

two  other  setae  caudad  of  spiracle;  at  caudal  margin  a  solitary  seta  close  to  ventral  margin. 
Sternum  with  scattered  slender  tubercles,  on  segment  3  these  appearing  as  a  longitudinal  row 
of  about  ten  alongside  the  hind  legs;  between  this  lateral  longitudinal  row  and  the  scattered 
discal  tubercles,  a  bare  space,  at  anterior  end  of  which  is  a  large,  stout,  setiferous  tubercle; 
a  subterminal  transverse  row  of  about  fifteen  acute  black  spines;  on  terminal  segments  the 
lateral  longitudinal  rows  interrupted,  or,  en  the  seventh  segment,  lacking,  on  seventh  segment 
the  subterminal  armature  of  both  dorsum  and  sternum  more  powerful,  especially  the  inter- 
mediate spines  of  sternum,  which  are  very  large;  numerous  setae  scattered  along  rows. 
Segments  3  to  8  on  ventral  surface  of  posterior  ring  with  a  circular  disklike  area,  median 
in  position  and  at  about  two-thirds  length  of  ring.  (In  some  specimens,  especially  females, 
the  discal  tubercles  on  the  posterior  ring  are  much  smaller  and  lie  in  longitudinal  rows.) 
Male  cauda  (Plate  XLV,  211)  with  tergal  valves  slender,  elongate,  slightly  divergent,  taper- 
ing to  the  acute  tips,  which  bear  a  small  subterminal  seta;  sternal  valves  short,  blunt,  with 
a  flattened  oval  lobe  between  valves  of  sternum;  eighth  segment  on  dorsum  with  a  trapezoid 
of  four  very  long,  slender,  pointed  lobes,  which  are  provided  with  delicate  hairs,  the  posterior 
pair  of  lobes  longer,  stouter,  and  lying  closer  together;  just  above  anterior  pair  of  lobes  a 
blunt  setiferous  tubercle;  pleural  region  of  segment  8  with  two  spines;  a  small  seta  cephalad 
of  these  spines;  sternal  region  of  segment  8  with  two  widely  separated  setiferous  tubercles. 
Female  cauda  (Plate  XLV,  212)  with  tergal  valves  elongate,  narrowed  at  tips,  and  armed 
with  sharp,  black  spines;  a  few  small  setae  before  tips;  sternal  valves  short,  blunt,  narrowed 
at  tips. 

Nepionotijpe. —  Orono,  Maine,  July  19,  1913. 
Neanotype. —  With  type  larva,  July  17,  1913. 

Paratypes. —  Larvae,  with  type  larva,  June  13  to  July  27, 1913.     Pupae,  June  9  to  July  22, 
1913. 

(Subgenus  Phylidorea  Bigot) 

1854     Phylidorea  Bigot.    Ann.  Soc.  Ent.  France,  p.  456. 

The  immature  stages  of  the  type  species  of  Phylidorea,  Limnophila 
(Phylidorea).  ferruginea  (Meig.),  were  described  by  De  Meijere  (1916: 
204-206).  In  Holland,  larvae  were  found  in  April  on  the  banks  of  a  ditch, 
where  they  were  living  between  decaying  leaves.  The  larvae  are  cylin- 
drical, and  are  from  brownish  yellow  to  brown  in  color.  The  antennae 
are  described  as  three-segmented,  the  apical  papilla  being  considered 
as  a  segment.  The  labrum  bears  on  the  epipharyngeal  surface  a  median 
projection  provided  with  two  conspicuous  biarticulate  lobes.  The  outer 
lobes  of  the  maxillae  project  as  flattened  blades.  The  mandibles  are 
powerful  and  strongly  curved,  and  have  a  group  of  small  lateral  teeth 
at  about  midlength. 

The  pupa  is  about  10  millimeters  long  and  is  blackish  brown  in  color. 
The  pronotal  breathing  horns  are  almost  semicircular  and  are  yellowish 


THE  CKANE-FLIES  OF  NEW  YORK  —  PART  II  867 

brown  in  color.  The  abdomen  is  depressed,  with  sharp  lateral  margins. 
The  abdominal  armature  is  described  as  consisting  of  small  hairs  instead 
of  slender  spines  as  in  the  American  species  of  this  subgenus. 

In  the  Nearctic  fauna,  the  subgenus  Phylidorea  includes  Limnophila 
adusta,  L.  similis  Alex.,  L.  novae-angliae  Alex.,  L.  lutea  Doane,  L.  terrae- 
novae  Alex.,  L.  costata  Coq.,  L.  fulvocostalis  Coq.,  L.  insularis  Johns.,  and 
probably  other  western  species.  It  seems  quite  possible,  moreover,  that 
the  species  allied  to  Limnophila  lento,  O.  S.  also  belong  here. 

Limnophila  (Phylidorea)  adusta  0.  S.  (supposition) 

1859    Limnophila  adusta  0.  S.    Proc.  Acad.  Nat.  Sci.  Phila.,  p.  235. 

A  larva  of  a  crane-fly  that  is  almost  certainly  a  member  of  this  genus 
was  taken  at  Ithaca,  New  York,  on  May  30,  1917,  in  company  with  a 
Ulomorpha  (No.  88-1917).  An  unknown  pupa  was  taken  in  the  sedge 
association  on  the  Bool  hillside,  Ithaca,  on  June  4,  1917.  It  was  not 
reared  and  its  identity  can  only  be  surmised,  but  there  can  be  little  doubt 
of  its  relationship  with  Phylidorea. 

Larva. —  Length,  8.6-9  mm. 
Diameter,  0.8  mm. 

Coloration  a  deep  saturated  orange-yellow,  the  body  with  silky  iridescent  reflections. 

Body  a  little  narrowed  at  both  ends.  Integument  with  a  long,  appressed,  golden 
pubescence.  Numerous  long  setae  and  pencils  of  hairs  on  body,  usually  at  about  midlength 
of,  or  on  posterior  half  of,  the  segments;  posterior  margin  of  each  segment  elevated  into  a 
prominent  transverse  ridge  of  hairs.  Spiracular  disk  surrounded  by  four  lobes,  ventral 
pair  about  twice  length  of  lateral  pair;  ventral  lobes  with  a  rather  narrow,  dark  brown  line 
on  inner  face;  outer  margin  fringed  with  very  long  dark  hairs  which  are  somewhat  paler 
at  their  bases,  this  fringe  of  hairs  longest  at  tips  of  lobes,  where  it  reaches  a  length  of  about 
eight  or  ten  times  length  of  lobe;  hairs  shorter  toward  base  of  lobe;  a  stiff  sensory  br'stle 
on  outer  face  of  lobe,  rather  far  back  from  tip;  lateral  lobes  similar,  with  the  fringe  of  hairs 
yellow  and  about  twice  length  of  lobes.  Spiracles  of  medium  size,  directed  toward  each  other. 
Anal  gills  four,  fleshy,  pale  in  color,  posterior  pair  longer  than  anterior  pair. 

Head  capsule  of  hexatomine  type,  the  dorsal  plates  narrowly  interrupted  on  mid-dorsal 
line  behind  clypeal-labral  sclerite.  Labrum  very  large,  anterior  margin  provided  with 
eight  or  ten  papillae  and  setae.  Epipharyngeal  region  having  a  large,  median,  circular 
area  which  is  slightly  elevated  and  bears  two  bisegmented  cylindrical  papillae.  Mental 
region  as  usual  in  the  genus,  consisting  of  three  hinged  bars  forming  three  sides  of  a  rectangle, 
the  anterior  transverse  bar  delicately  grooved.  Antenna  two-segmented,  second  segment 
narrower  than  first  and  bearing  at  its  tip  an  elongate  sculptured  papilla  and  a  long  seta. 
Mandible  of  hexatomine  type,  a  powerful  curved  hook,  at  about  midlength  of  which  is  an 


868  CHARLES  PAUL  ALEXANDER 

acute  tooth;  in  axil  of  latter  a  small  flattened  blade  which  is  slightly  widened  outwardly  and 
has  the  tip  truncated.     Maxillary  lobe  broad  at  base,  tapering  to  narrow  tip. 

Pupa. —  Length,  12  mm. 

Depth,  d.-v.,  1.6  mm. 

Similar  to  Limnophila  macrocera  in  general  shape  and  color,  differing  as  follows: 
Labrum  broadly  obtuse  at  tip.  Pronotal  breathing  horns  (Plate  XLIII,  203)  very  short, 
stout  but  flattened,  constricted  beyond  midlength,  and  with  a  row  of  breathing  pores  around 
apex;  pronotum  and  mesonotum carinate  medially  in  front;  wing  pad  showing  venation  rather 
clearly,  cell  Mi  deep,  basal  deflection  of  Cm  beyond  midlength  of  cell  1st  Mz-  Abdominal 
segments  with  lateral  margins  very  deeply  incised,  the  carinate  lateral  margins  very  accentu- 
ated, appearing  as  thin,  flattened  wings  (Plate  XLIII,  204).  Armature  of  abdomen  consist- 
ing of  abundant  elongate,  acicular  spines,  some  of  which  are  sinuously  twisted;  on  basal  ring 
these  spines  appearing  as  a  subterminal  transverse  row  on  both  dorsum  and  sternum;  on 
posterior  ring,  besides  the  subterminal  transverse  row,  spines  are  scattered  over  surface  in 
more  or  less  distinct  longitudinal  rows;  at  lateral  carina  a  group  of  about  a  dozen  long  spines 
at  caudal  margin,  as  well  as  a  powerful  spine  below  level  of  spiracle  on  extreme  margin  of 
carina;  ventrad  and  caudad  of  this,  three  long  setae,  two  close  beneath  spine  and  the  third 
underneath  spiracle.  Female  cauda  (Plate  XLIII,  202)  about  as  in  Limnophila  macrocera, 
but  sternal  valves  divergent  at  their  tips;  eighth  segment  with  a  trapezoid  of  dorsal  lobes 
about  as  in  L.  macrocera,  but  sternum  with  four  stout  spines  bearing  setae  on  their  sides; 
a  blunt  median  lobe  near  base  of  eighth  sternite. 

(Described  from  a  female  pupa  taken  on  Bool's  hillside,  Ithaca,  New  York,  on  June  4, 
1917,  where  it  was  associated  with  a  characteristic  swamp-inhabiting  crane-fly  fauna  — 
Bittacomorpha  clavipes,  Pseudolimnophila  luteipennis,  Limnophila  macrocera,  Pilaria  recondita, 
and  Prionocera  fuscipennis.) 

(Subgenus  Limnophila  Macquart) 

1834     Limnophila  Macq.     Suit.  &  Buff.,  vol.  1,  Hist.  Nat.  Ins.,  Dipt.,  p.  95. 
1863     Poecilostola  Schin.     Dipt.  Austriaca,  vol.  2,  p.  551. 

Limnophila  is  the  typical  subgenus  of  the  group,  the  type  having  been 
designated  as  Limnophila  pictipennis  by  Westwood  in  1840.  No  species 
of  this  subgenus  have  yet  been  described  from  North  America. 

The  larva  of  L.  punctata  (Meig.)  was  described  by  Beling  (1886:195- 
197)  and  by  Gerbig  (1913:158-161)  as  living  in  wet  earth,  by  Scheffer 
(1848: 10)  and  by  Cameron  (1917:63)  as  living  in  decaying  wood.  Gerbig 
found  larvae  in  sandy  soil  near  both  standing  and  flowing  water.  The 
larvae  are  very  active,  are  rust-brown  in  color,  and  attain  a  length  of  15 
millimeters  with  a  diameter  of  from  1.5  to  2  millimeters.  The  body  is 
provided  with  several  setae  and  setiferous  projections.  The  spiracular 
disk  (Plate  XL VI,  214)  is  surrounded  by  four  subequal  lobes  and  an  addi- 
tional reduced  dorso-median  lobe;  all  of  these  lobes  are  fringed  with  long 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  869 

hairs,  and  each  of  the  four  paired  lobes  has  a  sensory  bristle  near  the  tip. 
The  four  anal  gills  are  not  very  prominent. 

The  head  capsule  is  of  the  hexatomine  type.  The  mandibles  are 
sickle-shaped,  with  two  small,  broad,  blunt,  sawlike  teeth  just  beyond 
midlength. 

Brauer  (1883:55)  found  the  larvae  of  another  species  —  L.  pictipennis 
(Meig.),  the  type  of  the  genus  —  between  wet  decaying  leaves  in  bogs. 
His  figure  of  the  head  capsule  (1883,  pi.  1,  fig.  10)  shows  a  typical  hexa- 
tomine head  (Plate  XL VI,  213).  Beling  (1879:51-52).  found  this  species 
in  the  sand  of  a  small,  dried-out  brook  bed. 

i 
Group  Ulomorphae 

Genus  Ulomorpha  Osten  Sacken  (Gr.  Ula  +  shape)  \ 

1869      Ulomorpha  0.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  232. 

Larva.—  Form  slender.  Body  covered  with  a  rich  golden-yellow  pubescence.  Spiracular 
disk  surrounded  by  four  unequal  lobes,  the  ventral  pair  the  longest,  lying  subparallel,  fringed 
with  exceedingly  elongate  hairs.  Spiracles  small.  Head  capsule  narrow,  the  dorsal  plate 
narrow,  at  tip  expanded  into  a  spatula.  Mandible  hinged,  blade  very  long  and  slender, 
with  a  single  very  long  lateral  tooth  at  its  base.  Maxilla  densely  golden  hairy,  outer  lobe 
projecting,  bladelike.  Antenna  with  a  very  long,  tapering,  apical  papilla.  Mentum  not 
chitinized. 

Pupa.—  Cephalic  crest  prominent,  each  lobe  with  three  setae.  Pronotal  breathing  horns 
very  long  and  slender,  sinuous,  cylindrical,  at  apex  split  into  two  flattened  divergent  lobes. 
Mesonotum  short,  very  convex,  unarmed.  Wing  sheaths  attaining  end  of  second  abdominal 
segment.  Leg  sheaths  short,  ending  before  tip  of  third  abdominal  segment;  hind  tarsi 
a  little  longer  than  the  others.  Abdominal  segments  divided  into  two  rings,  posterior  ring 
with  a  subterminal  transverse  armature  of  stout  black  spines  and  a  few  long  setae;  sternal 
armature  stronger  than  that  of  dorsum.  Pleura  armed  with  a  few  similar  spines.  Acido- 
thecae  of  ovipositor  very  long  and  slender.  Dorsum  of  segment  8  with  a  trapezoid  of  four 
lobes. 

The  genus  Ulomorpha  includes  five  known  species,  the  genotype, 
Ulomorpha  pilosella,  of  the  eastern  United  States,  and  four  western 
North  American  species.  The  immature  stages  of  the  genotype  are  spent 
in  rich  organic  mud  in  shaded  situations. 

Ulomorpha  pilosella  (O.  S.) 

1859    Limnophila  pilosella  0.  S.    Proc.  Acad.  Nat.  Sci.  Phila.,  p.  242.  " 
Ulomorpha  pilosella  is  not  uncommon  in  cool  Canadian  woods  thruout 
northeastern  North  America,   and  the  adult  flies  may  be  swept  from 


870  CHARLES  PAUL  ALEXANDER 

rank  vegetation  in  such  haunts.  The  flies  bear  a  marked  resemblance 
to  the  species  of  the  subgenus  Lasiomastix  of  the  genus  Limnophila,  but 
the  pupae,  especially  in  the  structure  of  the  pronotal  breathing  horns, 
are  quite  distinct.  The  larvae  are  different  from  the  related  and  some- 
what similar  larvae  of  the  subgenus  Dicranophragma  of  the  genus  Lim- 
nophila, of  Penthoptera,  and  of  similar  hexatomine  genera,  in  their  pale 
whitish  yellow  coloration  instead  of  the  deep  saturated  yellows  and  oranges 
of  the  genera  mentioned.  All,  however,  have  the  quick,  restless  move- 
ments so  characteristic  of  this  group  of  crane-flies.  Larvae  found  on 
Bool's  hillside,  Ithaca,  New  York,  on  May  14  and  23,  1917,  transformed 
to  adults  on  June  9. 

Larva. —  Length,  8.5-9  mm. 

Diameter,  0.5-0.6  mm. 

Coloration  pale  whitish  yellow,  the  eighth  abdominal  segment  suddenly  whitish. 

Form  slender.  Body  covered  with  a  dense  golden-yellow  pubescence.  A  transverse  fringe 
of  stiff,  erect,  short  hairs  at  posterior  margin  of  prothorax.  A  number  of  pencils  of  setae 
or  solitary  bristles  on  sides  of  segments,  one  on  each  annulus,  longest  near  posterior  margins 
of  segments.  Behind  these  setae,  tufts  of  small  hairs.  Spiracular  disk  (Plate  XLVI, 
219)  surrounded  by  four  lobes;  ventral  pair  the  longest,  lying  subparallel  to  each  other, 
outer  margin  fringed  with  long,  delicate  hairs,  those  near  tip  coarse  and  easily  broken,  some 
of  the  hairs  at  tips  exceedingly  elongate;  inner  face  of  ventral  lobes  heavily  suffused  with  dark 
brown,  this  color  more  intense  proximally;  lateral  lobes  short,  with  an  apical  fringe  of  coarse 
yellowish  setae.  Spiracles  very  small,  widely  separated,  located  at  base  of  lateral  lobes. 
Anal  gills  four,  very  slender,  pale  in  color,  the  posterior  pair  a  little  the  longer.  On  sternum 
of  eighth  abdominal  segment,  before  gills,  a  transverse  row  of  four  long,  coarse  setae. 

Head  capsule  (Plate  XLVI,  215)  very  long  and  narrow,  the  dorsal  plate  slender,  at  end 
expanded  into  a  spatula;  lateral  plates  a  little  shorter  than  dorsal  plate.  Labrum  (Plate 
XLVI,  216)  and  epipharynx  broadly  transverse,  projecting,  the  anterior  margin  narrower, 
truncated,  on  either  side  near  base  with  a  brush  of  long  hairs;  disk  of  epipharyngeal  region 
with  four  setae,  posterior  pair  a  little  the  closer  together;  a  few  tiny  papillae  on  ventral 
surface;  clypeal  region  emarginate,  with  two  large  setae  near  anterior  margin  and  another 
immediately  behind  base  of  antenna.  Mental  region  not  readily  distinguishable  in  the 
material  available,  but  at  the  most  with  little  or  no  chitinization.  Antenna  (Plate  XLVI, 

217)  with  basal  segment  cylindrical,  a  little  narrowed  medially,  the  truncated  apex  with  about 
two  or , three  long  setae  and  a  very  long,  hyaline,  sensory  papilla  which  tapers  gradually  to 
apex,  this  papilla  about  three  times  length  of  segment  bearing  it.     Mandible  (Plate  XLVI, 

218)  hinged,  the  base  slender  but  powerful,  with  the  inner  face  deeply  concave  to  receive 
mandible  in  a  position  of  rest;  blade  of  mandible  produced  into  a  very  slender  hook  which 
is  almost  straight,  a  little  curved  at  extreme  tip,  at  its  base  a  very  large,  acute,  flattened 
blade  which  is  more  than  half  length  of  mandible  itself;  in  its  angle  this  blade  has  a  second, 
microscopic,  tooth;  prostheca  with  about  five  long,  stout,  comblike  teeth  exceeding  the 
mandible  in  length,  and  an  additional  shorter,  flattened  blade  marked  with  parallel  grooved 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  871 

lines.  Maxilla  with  dense  tufts  of  long  yellow  hairs;  outer  lobe  produced  cephalad  as  a 
hyaline,  flattened  blade  which  projects  from  prothoracic  orifice  when  head  is  retracted. 

Pupa. —  Length,  8.5-9  mm. 

Length  of  pronotal  breathing  horns,  nearly  2  mm. 
Width,  d.-s.,  1  mm. 
Depth,  d.-v.,  1  mm. 

/  Head,  thorax,  and  appendages  dark  brown;  pronotal  breathing  horns  similar,  but  terminal 
half  gradually  paler,  the  tip  almost  yellow;  abdomen  pale  brown. 

Cephalic  crest  consisting  of  two  prominent  lobes,  each  with  three  strong  setae,  the  most 
ventral  directed  outward;  just  before  primary  crest  and  lying  between  antennal  bases,  a  very 
low,  slightly  bilobed  crest  which  is  not  setiferous.  Labrum  elongate,  obtusely  rounded  at 
apex  and  separating  labial  lobes,  the  latter  produced  caudally  into  subacute  points.  Maxillary 
palpi  stout  at  base,  narrowed  to  tip  (Plate  XLVII,  221).  Antenna  short  (in  female  sex,  at 
least),  extending  but  a  short  distance  beyond  knee  joints  of  fore  legs.  Pronotoim  (Plate 
XLVII,  220)  high,  feebly  carinate  medially.  Breathing  horns  separated  basally,  very  long 
and  slender,  sinuous,  transversely  wrinkled,  at  tip  split  into  two  flattened  divergent  lobes 
(Plate  XLVII,  222  and  223).  Mesonotum  very  short  and  convex,  with  numerous  black  dots 
which  are  most  abundant  anteriorly.  Two  small  setae  on  either  side  behind  wing  axilla. 
Lateral  angles  of  mesonotum  blunt,  but  tip  produced  into  a  slender  setiferous  tubercle. 
Wing  sheaths  ending  opposite  tip  of  second  abdominal  segment.  Leg  sheaths  short,  ending 
just  before  tip  of  third  abdominal  segment,  hind  legs  a  little  longer  than  others. 

Abdomen  with  a  narrow  basal  ring  and  a  much  broader  posterior  ring,  the  latter  armed 
before  posterior  margin  with  a  transverse  row  of  small  black  spines,  strongest  on  pleura, 
weakest  on  dorsum;  on  dorsum  (Plate  XLVII,  224)  the  spines  reduced  in  number,  there 
being  from  one  to  five  (or  in  some  cases  none),  and  occurring  only  at  or  near  ends  of  row; 
ends  of  row  with  two  setae;  on  either  side  of  median  line  a  group  of  three  closely  approxi- 
mated setae;  usually  segments  2  and  3  have  the  spines  weak  or  lacking;  segments  4  to  6 
with  two  spines,  and  segment  7  with  one  spine,  but  in  some  specimens  the  number  is  slightly 
increased.  Sternites  (Plate  XLVII,  225)  with  the  intermediate  segments  (4  to  6)  having 
about  twenty  spines  in  an  almost  continuous  row  which  as  a  rule  is  uninterrupted;  at 
each  end  of  row  about  two  strong  setae;  near  base  of  posterior  ring  a  narrow  transverse 
area  with  two  setae  at  each  end.  Pleura  with  a  few  powerful  spines,  small  or  lacking  on 
basal  segments,  larger  and  more  numerous  on  posterior  segments,  there  being  usually  two 
on  segment  4,  three  on  segments  5  and  6,  and  four  on  segment  7;  on  pleura  at  about  mid- 
length  of  posterior  ring  and  nearer  dorsal  side,  three  black  setae  in  transverse  alinement, 
these  somewhat  longer  on  basal  segments;  opposite  basal  ring  a  stout  seta.  Female  cauda 
(Plate  XLVII,  228)  elongate;  tergal  valves  slender,  slightly  upcurved,  near  apex  with  a 
sharp  black  spine  which  is  directed  dorsad,  laterad,  and  caudad;  two  weak  setae  on  either 
side  before  apex.  Dorsum  of  segment  8  with  four  lobes;  posterior  pair  elongate,  slender, 
curved,  and  divergent;  anterior  pair  blunt,  small,  and  more  approximated;  just  ventrad  of 
these  lobes  a  stout  seta;  pleural  region  with  two  powerful  spines,  above  the  more  dorsal 
of  which  is  a  stout  seta;  a  seta  near  ventral  margin. 

Nepionotype.—  Ithaca,  New  York,  May  30,  1917.     No.  88-1917. 
Neanotype. —  With  type  larva,  reared  June  9,  1917. 
Paratypes  —  Larvae,  May  30,  1917.     Pupa,  June  13,  1917. 


872  CHARLES  PAUL  ALEXANDER 

Genus  Pilaria  Sintenis  (Lat.,  derived  from  the  long  antennal  verticils) 

1888     Pilaria  Sintenis.     Sitzber.  Nat.-Ges.  Dorpat.,  vol.  8,  p.  398. 

1919     Eulimnophila  Alex.     Cornell  Univ.  Agr.  Exp.  Sta.,  Mem.  25,  p.  917. 

Larva.- — Form  moderately  slender.  Spiracular  disk  surrounded  by  four  unequal  lobes, 
lateral  pair  in  some  cases  very  reduced,  elongate  ventral  lobes  fringed  with  long  hairs.  Head 
capsule  of  Ulomorpha  type.  Mandible  hinged,  blade  with  one  or  two  acute  teeth  at  base. 
Maxilla  densely  hairy.  Mentum  not  chitinized. 

Pupa. —  Pronotal  breathing  horns  elongate-cylindrical,  tips  split  into  flattened  lobes. 
Abdominal  segments  with  three  or  four  pairs  of  naked  tubercles. 

The  genus  Pilaria  includes  a  group  of  species  of  the  old  genus  Limnophila, 
comprising  Limnophila  tenuipes  and  L.  pilicornis  and  their  allies.  Its 
relationships  are  plainly  with  Ulomorpha  rather  than  with  Limnophila. 
The  genotype  is  Limnophila  pilicornis  (Zett.),  of  northern  Europe.  Other 
included  species  are  L.  tenuipes,  L.  recondita,  L.  imbecilla  O.  S. ,  L.  edwardi 
Alex.,  L.  quadrata,  L.  stanwoodae  Alex.,  and  L.  osborni  Alex.,  of  North 
America,  and  L.  discicollis  (Meig.),  L.  fuscipennis  (Meig.),  L.  subtincta 
(Zett.),  and  probably  other  species,  of  Europe. 

The  immature  stages  are  spent  in  mud  or  moist  earth.  In  Europe, 
Pilaria  discicollis  (Plate  XL VIII,  232)  has  been  found  by  Gerbig  (1913: 
163-164)  and  by  Cameron  (1917:63).  P.  fuscipennis  is  described  by 
Beling  (1886 : 197-198)  as  living  in  mud  near  a  ditch.  Gerbig  (1913 : 164- 
166)  found  the  larvae  (Plate  XL VIII,  231)  in  a  similar  situation.  Brauer 
(1883:54)  describes  them  as  living  between  decaying  leaves  in  swamps. 
Cameron  (1917:63)  states  that  the  larvae  are  found  in  decaying  wood, 
this  record  possibly  being  an  error.  According  to  Beling,  the  pupal  dura- 
tion is  not  more  than  ten  days.  In  America,  P.  tenuipes  has  been  dis- 
cussed by  Hart  (1898  [1895]: 204-205)  and  by  Malloch  (1915-17b:223- 
224),  as  stated  under  the  discussion  of  the  species. 

The  species  of  the  genus  Pilaria  may  be  separated  by  the  following 
keys: 

Larvae 

Ventral  lobes  of  spiracular  disk  elongate,  heavily  marked  with  brownish  black;  coloration 

pale  yellow P.  tenuipes  (Say)  (p.  873) 

Ventral  lobes  of  spiracular  disk  short,  pale;  coloration  deep  yellow. 

P.  recondita  (O.  S.)  (p.  874) 
Pupae 

1.  Pronotal  breathing  horns  short,  black;  lobules  of  cephalic  crest  blunt  and  rounded. 

P.  quadrata  (O.  S.)  (p.  875) 

Pronotal  breathing  horns  elongate,  pale,  brownish  yellow  or  yellow;  lobules  of  cephalic 
crest  elongate,  finger-like 2 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  873 

2.  Antenna!  sheaths  of  male  elongate;  pronotal  breathing  horns  longer,  pale  yellow. 

P.  tenuipes  (Say)  (p.  873) 
Antennal  sheaths  of  male  short;  pronotal  breathing  horns  shorter,  yellowish  brown. 

P.  recondita  (O.  S.)  (p.  874) 

Pilaria  tenuipes  (Say) 

1823    Limnobia  tenuipes  Say.     Journ.  Acad.  Nat.  Sci.  Phila.,  vol.  3,  p.  21. 
1869    Limnophila  tenuipes  O.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  210-211. 

Pilaria  tenuipes  is  a  widely  distributed  crane-fly  thruout  eastern  North 
America.  The  immature  stages  are  commonly  found  in  the  mud  of 
swamps,  or  near  streams  and  other  bodies  of  water.  This  is  the  unknown 
Limnophila  described  by  Hart  (1898  [1895]:  204-205),  and  also  considered 
in  much  detail  by  Malloch  (1915-17  b:  223-224),  who  found  the  pupae 
along  the  banks  of  the  Sangamon  River  in  Illinois. 

Larva. —  Length,  16  mm. 

Diameter,  1.4-1.5  mm. 

Coloration  of  living  larva,  pale  brownish  yellow.  Body  covered  with  a  long,  dark-colored, 
appressed  pubescence,  more  conspicuous  on  posterior  segments.  Lateral  pencils  of  setae 
near  base  and  apex  of  segments.  Antepenultimate  segment  of  body  capable  of  globular 
distention,  covered  with  numerous  transverse  rows  of  microscopic  roughened  points. 
Spiracular  disk  (Plate  XLVIII,  230)  moderately  large,  surrounded  by  four  lobes;  ventral 
lobes  long  and  slender,  inner  face  with  closely  approximated,  transverse,  brownish  black  lines 
which  cause  entire  face  to  appear  dark;  near  tips  of  lobes  these  black  marks  tapering  out 
into  a  long  point;  basal  parts  of  dark  marks  subcontiguously  hollowed  out  interiorly  to  form 
a  large  pale  area  below  spiracles;  ventral  lobes  fringed  with  long,  pale  hairs,  some  of  them 
exceedingly  elongate.  Lateral  lobes  very  small,  blunt,  tending  to  be  reduced,  bearing  short 
fringes  of  dark  hairs.  Anal  gills  slender,  pale  in  color. 

Head  capsule  of  Ulomorpha  type  and  not  very  different  from  that  of  the  type  genus; 
dorsal  plate  broad  basally,  narrowed  gradually  behind  to  near  tip  where  it  expands  into 
a  very  large  spatula.  Epipharyngeal  region  of  labrum  and  maxillae  fringed  with  dense  tufts 
and  brushes  of  long  yellow  hairs.  Maxillary  lobe  relatively  small  but  elongate,  hyaline, 
tapering  to  flattened  apex.  Antenna  (Plate  XLVIII,  229)  with  basal  segment  elongate, 
bearing  at  its  tip  an  elongate  apical  papilla  which  is  a  little  longer  than  the  segment,  bluntly 
rounded  at  its  tip,  and  delicately  sculptured.  Mandible  (Plate  XLVIII,  228)  very  long, 
hinged,  at  its  base  an  acute  tooth  equal  in  length  to  about  one-third  length  of  mandible;  in 
the  type  larva,  the  left  mandible  a  little  longer  than  the  right  mandible.  Mental  region 
not  chitinized. 

Pupa. —  Length,  10-15  mm. 

Width,  d.-s.,  1.5-1.6  mm. 
Depth,  d.-v.,  1.6-1.8  mm. 

Coloration  dark  brown;  pronotal  breathing  horns  light  yellow,  extreme  bases  brownish. 

Cephalic  crest  small,  composed  of  three  slender,  finger-like  lobes  which  are  tipped  with 

strong  setae;  on  front,  before  crest,  two  setiferous  lobes.     Labrum  small,  bluntly  rounded 


874  CHARLES  PAUL  ALEXANDER 

at  apex.  Labial  lobes  oval.  Maxillary  palpi  slender.  Antenna  of  female  moderately 
elongated,  reaching  to  just  beyond  wing  root.  Pronotal  breathing  horns  (Plate  XLIX,  233) 
elongate,  cylindrical,  sinuous,  transversely  wrinkled,  apex  scarcely  enlarged  but  deeply 
split  on  inner  margin;  a  setiferous  tubercle  ventrad  and  laterad  of  breathing  horns.  Thorax 
with  a  very  high  anterior  median  crest.  A  slender,  setiferous  tubercle  above  wing  axil, 
and  two  others  on  either  side  of  median  line.  Sheaths  of  halterss  long  and  slender.  Leg 
sheaths  ending  about  on  a  level,  or  those  of  hind  legs  the  shortest  and  those  of  fore  legs  a 
little  longer. 

Abdominal  segments  divided  into  two  anmili.  Tergites  (Plate  XLIX,  235)  on  posterior 
ring  with  a  caudal  row  of  blunt,  naked  tubercles;  at  end  of  row  two  setae;  on  either  side  of 
median  area  and  just  in  front  of  row,  a  large  setiferous  tubercle;  at  base  of  ring  two  naked 
tubercles,  one  on  either  side  of  median  line;  two  slender  setiferous  tubercles  near  margin 
of  ring.  Basal  ring  with  four  naked  tubercles.  (The  third  pair  of  tubercles  found  in  the 
pupa  of  Pilaria  quadrata  is  vestigial.)  Pleura  with  four  tubercles,  two  on  each  ring,  the 
basal  one  of  each  ring  setiferous,  the  posterior  one  naked.  Sternites  with  six  naked  tubercles 
on  basal  ring,  arranged  in  three  transverse  pairs;  on  posterior  ring  at  base  two  setiferous 
tubercles,  each  with  two  bristles,  directly  behind  last  naked  tubercle  of  basal  ring;  at  caudal 
margin  two  or  three  large  tubercles  near  end  of  row  and  about  four  or  five  small  naked  tubercles 
between.  Female  cauda  (Plate  XLIX,  234  and  236)  very  elongate,  tergal  valves  slightly 
upcurved,  terminating  in  a  sharp  spine  and  with  two  setae  on  outer  face.  Eighth  tergite 
with  four  elongate  lobes;  posterior  pair  blunt,  directed  laterad;  anterior  pair  elongate,  slender, 
with  two  setae  laterad  of  each.  A  blunt  lobe  on  pleural  region.  Pleura  with  three  or  four 
powerful  tubercles,  with  a  seta  located  between  the  more  dorsal  pair.  Sternal  region  with 
four  small  setae,  two  on  either  side  of  broad  median  area. 

Nepionotype. —  Orono,  Maine,  July  1,  1913.     No.  50-1913. 
Neanotype. —  With  type  larva. 

Pilaria  recondita  (O.  S.) 

1869    Limnophila  recondita  0.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  212-213. 

Pilaria  recondita  is  a  common  crane-fly  thruout  the  northeastern 
United  States.  The  immature  stages  are  swamp  inhabitants,  and  are 
very  frequently  found  in  exactly  the  same  situations  as  are  those  of  P. 
tenuipes.  P.  recondita  belongs  to  the  same  group  as  P.  tenuipes,  and  the 
pupae  of  the  two  species  are  very  difficult  to  distinguish. 

Larva. —  Length,  15  mm. 

Diameter,  1.2  mm. 

Color  a  uniform  light  yellow. 

Body  covered  with  a  long,  appressed,  yellow  pubescence  and  with  a  few  long  setae. 
Spiracular  disk  very  small,  in  a  position  of  rest  almost  closed,  surrounded  by  four  lobes; 
ventral  lobes  moderately  elongated,  fringed  with  long,  golden-yellow  hairs  which  are  longest 
at  tips  of  lobes;  if  bent  backward  these  elongate  hairs  extending  to  beyond  gills;  inner  face 
of  lobes  almost  unmarked,  with  only  a  delicate  brown  line  extending  from  tip  toward  base 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  875 

for  a  distance  equal  to  about  one-half  length  of  lobes;  lateral  lobes  small,  subdorsal  in  position, 
separated  by  a  narrow  notch,  their  inner  faces  opposed  to  each  other,  margin  fringed  with 
short,  golden-yellow  hairs.  Anal  gills  four,  moderately  elongated. 

Head  capsule  as  in  P.  tenuipes.  Antenna  with  sculptured  apical  papilla  tapering  to 
blunt  tip;  besides  this  papilla,  an  even  longer,  hyaline,  flattened  blade.  Mandible  with 
apical  bladelike  part  shorter  and  stouter,  with  two  subequal  stout  triangular  teeth  at  base 
(Plate  XLVIII,  227). 

Pupa.' — Very  similar  to  pupa  of  P.  tenuipes,  but  smaller.  Antennal  sheaths  of  male 
short.  Breathing  horns  a  little  shorter  than  in  P.  tenuipes  but  still  much  longer  than 
in  P.  quadrata,  of  a  pale  yellowish  brown  color.  On  abdominal  tergites,  along  caudal  margin 
of  posterior  ring,  from  four  to  seven  naked  tubercles  between  the  setiferous  tubercles  (in 
P.  tenuipes,  four  or  five).  Male  cau da.  (Plate  L,  237)  with  dorsal  lobes  stout,  cylindrical, 
narrowed  at  tips,  divergent,  direstsd  caudad  and  ventrad;  on  outer  face  before  tip  a  slender 
seta;  ventral  lobes  blunt,  with  a  flattened  ventral  tubercle  at  base 'of  notch.  Segments  on 
dorsum  with  posterior  lobes  blunt,  straight,  directed  caudad  and  slightly  laterad,  but  not  so 
strongly  as  in  P.  tenuipes. 

Nepionotype. —  Orono,  Maine,  July  3,  1913. 

Neanotype  —  Ithaca,  New  York,  emerged  June  11,  1917.     No.  112-1917. 

Paratypes. —  Pupa,  Orono,  Maine,  placed  in  rearing  as  a  fully  grown  larva,  June  26,  1913; 
emerged  as  an  adult  male,  July  3,  1913,  showing  a  pupal  duration  of  seven  days.  Larva, 
Orono,  Maine,  July  5,  1913  (No.  74-913). 

Pilaria  quadrata  (O.  S.) 

1859    Limnophila  quadrata  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  241. 

Pilaria  quadrata  is  a  widely  distributed  spring  and  early  summer 
species.  The  immature  stages  are  very  similar  to  those  of  P.  tenuipes 
and  P.  recondita.  A  pupa  was  found  by  Dr.  Needham  in  the  Indian 
Spring,  Ithaca,  New  York,  where  it  was  found  floating  among  the  water 
cress.  From  this  pupa  an  adult  female  fly  was  reared.  On  June  3,  1917, 
the  writer  found  two  fully  matured  male  pupae  in  Chickaree  Woods  near 
Ithaca.  There  had  been  a  very  heavy  rainstorm  on  the  preceding  day, 
and  the  low  spots  in  the  woods  had  been  converted  into  small  ponds, 
many  of  the  insects  that  normally  live  in  the  mud  or  beneath  the  decaying 
leaves  being  forced  to  the  surface.  The  pupae  of  P.  quadrata,  as  well 
as  an  abundance  of  Tipula  larvae,  were  found  clinging  to  small  islands 
of  debris  floating  on  these  temporary  woodland  pools.  The  adult  flies 
emerged  on  June  3. 

Pupa.—  Length  of  cast  pupal  skin,  9-12  mm. 

Coloration  almost  black,  including  pronotal  breathing  horns;  abdomen  more  dusky  gray. 

Cephalic  crest  small,  black,  trilobed,  each  lobe  with  a  seta  at  apex.     Labrum  narrow, 

blunt  at  tip.     Labial  lobes  rounded.     Sheaths  of  maxillary  palpi  elongate,  tapering  to  the 


876  CHARLES  PAUL  ALEXANDER 

slender  points.  Antenna  short  in  both  sexes.  Pronotal  breathing  horns  (Plate  L,  238) 
moderately  elongate,  cylindrical,  transversely  wrinkled,  at  tips  smooth,  flattened,  and 
slightly  enlarged.  A  tubercle  with  two  long  setae  above  wing  axil.  Two  satiferous  punctures 
on  dorsum  on  either  side  of  median  line.  Wing  sheaths  ending  before  tip  of  second  abdominal 
segment.  Leg  sheaths  ending  before  tip  of  third  abdominal  segment,  the  tarsal  sheaths 
ending  about  on  a  level  or  the  hind  legs  shorter. 

Abdominal  segments  with  tergites  (Plate  L,  239)  2  to  6  provided  with  eight  naked, 
discal  tubercles,  arranged  in  four  transverse  pairs,  the  third  pair  more  approximated;  laterad 
of  third  pair  of  naked  tubercles,  two  small  setiferous  tubercles;  on  seventh  segment  one  of  the 
four  pairs  of  tubercles  lacking;  near  caudal  margin  of  segments  a  transverse  row  of  weak, 
setiferous  tubercles;  on  either  side  of  median  line,  in  alinement  with  discal  tubercles  and 
just  anterior  to  the  transverse  setiferous  row,  a  large  tubercle  provided  with  three  setae. 
Pleural  region  carinate,  each  segment  armed  with  four  slightly  curved  tubercles:  anterior 
one  solitary,  setiferous;  sacond  one  solitary,  nakad;  third  one  with  two  or  three  setae;  posterior 
one  bifid,  naked.  Sternites  (Plate  L,  240)  with  six  naked,  discal  tubercles  corresponding 
to  those  of  tergites  but  reduced  in  number.  Subterminal  armature  weak,  ends  of  rows 
tuberculate;  an  isolated  setiferous  tubercle  ventrad  and  laterad  of  ends  of  rows.  Male 
cauda  (Plate  L,  242)  elongate;  dorsal  lobes  elongate-cylindrical,  directed  caudad  and 
slightly  dors^i,  tapering  to  acute  tips;  three  weak  setae  on  outer  ventral  face;  ventral  lobes 
blunt,  much  shorter  than  dorsal  lobes,  with  a  blunt  median  lobule  between  them  at  their 
base;  eighth  segment  on  tergum  provided  with  a  large,  blunt,  median  tubercle,  with  two 
large  posterior  lobes  which  are  directed  caudad  and  with  two  tiny  lobes  on  either  side  in 
front;  sternum  with  a  transverse  row  of  four  separated  setiferous  tubercles;  caudal  margin 
with  a  transverse  row  of  about  eight  or  nine  pale  tubercles  on  either  side,  the  outermost  the 
largest;  a  small  seta  above  second  tubercle  at  ends  of  row.  Female  cauda  (Plate  L, 
241)  very  long  and  slender,  subacicular,  sternal  valves  a  little  shorter  than  tergal  valves; 
tergal  valves  terminating  in  blunt  cylindrical  points. 

Neanotype. —  Male  pupal  skin,  Ithaca,  New  York,  June  3,  1917. 
Paratypes. —  Pupa,  a  male  skin  with  type  pupa;  a  female  skin,  Ithaca. 

Subtribe  Hexatomaria 

The  subtribe  Hexatomaria  comprises  a  well-defined  division  with  but 
four  known  genera,  three  of  which  are  North  American  and  are  considered 
in  this  paper.  The  only  other  group  of  crane-flies  with  which  the  species 
may  be  confused  are  certain  of  the  Limnophilaria,  especially  the  Ulomorpha 
group  of  genera. 

The  larvae  havo  the  labral  sclerite  of  the  head  capsule  large,  separated 
from  the  remainder  of  the  capsule  by  a  distinct  suture.  The  epipharyngeal 
region  is  restricted  to  the  anterior  median  part  of  the  sclerite,  and  is  pro- 
vided with  two  large  tubercles  on  either  side,  which  are  tipped  with  two 
or  ^.hree  hyaline,  cylindrical  papillae.  Between  these  papillae  is  a  brush 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  877 

of  hairs  surrounding  two  pairs  of  setiferous  tubercles.  The  mental  region 
is  not  chitinized.  The  dorsal  plates  of  the  head  capsule  are  widely  sepa- 
rated from  each  other  by  a  median  split. 

The  pupae  are  often  armed  with  spines  or  tubercles  on  or  about  the 
head  and  the  thorax.  The  lateral  abdominal  spiracles  are  large  and 
functional. 

The  genera  of  the  subtribe  Hexatomaria  may  be  separated  by  the 
following  keys: 

Larvae 

1.  Coloration  of  body  a  deep  saturated  orange-yellow;  spiracular  disk  with  ventral  lobes 

unlined  with  darker,  bearing  at  tips  a  few  very  long  hairs.  . .  Penthoptera  Schin.  (p.  891) 

Coloration  of  body  pale  yellow,  whitish,  or  greenish;  spiracular  disk  with  lobes  lined 

with  dark  brown  or  black 2 

2.  Size  small,  form  slender  (length  14-15  mm.,  diameter  1-1.3  mm.) .  .  Hexaloma  Latr.  (p.  877) 
Size  larger,  form  stouter  (length  over  15  mm.,  diameter  over  1.6  mm.) 

Eriocera  Macq.  (p.  881) 
Pupae 

1.  Pronotal  breathing  horns  short  and  stout,  red  at  base  and  apex,    the  remaining  part 

dark-colored,  transversely  wrinkled;  horns  bent  strongly  toward  each  other  at  tips. 

Penthoptera  Schin.  (p.  891) 
Pronotal  breathing  horns  not  as  above 2 

2.  Size  small  (length  under  10  mm.) Hexatoma  Latr.  (p.  877) 

Size  larger  (length  over  12  mm.) .- Eriocera  Macq.  (p.  881) 

Genus  Hexatoma  Latreille  (Gr.  six  +  /  cut) 

1809    Hexatoma  Latr.     Gen.  Crust,  et  Ins.,  vol.  4,  p.  260. 

1818     Nematocera  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  1,  p.  209. 

1818    Anisomera  Meig.    Syst.  Beschr.  Zweifl.  Ins.,  vol.  1,  p.  210. 

Larva. —  Size  small  (length  of  Hexatoma  megacera  about  14  mm.) .  Spiracular  disk  sur- 
rounded by  two  pairs  of  lobes,  ventral  pair  the  longer,  inner  face  marked  with  a  narrow 
brown  line  which  is  expanded  at  its  inner  end.  Head  capsule  with  lateral  angles  of  labrum 
elongate  and  densely  clothed  with  hairs. 

Pupa.—  Size  small  (length  under  10  mm.).  A  large  spinous  tubercle  on  scape  of  antenna. 
No  median  projection  on  mesonotal  scutellum.  Pronotal  breathing  horns  short  and  straight. 
Wing  pads  with  cell  jR2  very  small  and  M  with  but  a  single  branch  reaching  wing  margin. 

The  genus  Hexatoma  includes  a  small'  number  of  forms  with  a  chiefly 
Holarctic  distribution,  there  being  about  eleven  Palearctic,  two  Ethiopian, 
and  one  North  American  species  so  far  described.  The  adult  flies  of 
the  North  American  species,  Hexatoma  megacera,  are  common  on  vegetation 
along  the  banks  of  rather  large  streams.  They  are  discussed  herein  in 
detail  under  the  account  of  this  species.  The  adult  flies  of  some  of  the 
larger  European  species  have  habits  quite  like  those  of  the  genus  Eriocera, 


878  CHARLES  PAUL  ALEXANDER 

as  discussed  elsewhere  in  this  paper  (page  705).  Kiedel  (1909:29) 
describes  in  some  detail  the  habits  and  swarming  of  H.  bicolor  (Meig.), 
the  males  of  which  are  very  active  in  the  forenoon  during  the  hours  of 
brightest  sunlight,  the  females  resting  on  the  willow  branches  near  by. 
Similar  habits  are  recorded  by  Riedel  (1910:30)  for  H.  saxonum  (Lw.). 
The  immature  stages  of  the  European  species  are  practically  unknown, 
the  only  original  reference  being  that  of  Von  Roser  (1834),  who  states  that 
the  larvae  of  H.  nigra  Latr.  live  in  the  sand  along  the  banks  of  streams. 

Hexatoma  megacera  (0.  S.) 

1859    Anisomera  mejzcera  0.  S.    Proc.  Acad.  Nat.  Sci.  Phila.,  p.  242. 

The  adult  flies  of  Hexatoma  megacera  are  on  the  wing  during  the  months 
of  May  and  June  and  may  be  swept  from  the  rank  vegetation  along  the 
streams  from  which  their  larvae  emerged.  The  following  notes  on  copu- 
lation, resting  positions,  egg  laying  in  nature  and  in  captivity,  and  other 
details,  are  quoted  from  published  field  observations  (Alexander,  1915  c: 
143-145) : 

May  14,  1911  —  This  usually  rare  insect  was  common  on  a  grassy  plot  of  land  along 
Cascadilla  Creek  [Ithaca,  New  York].  The  flies  sit  on  the  blades  of  grass,  the  long  antennae 
of  the  male  directed  straight  ahead.  The  males  are  very  poor  fliers  and  prefer  to  drop  to  the 
ground  when  disturbed  and  clumsily  work  their  way  off  along  the  ground.  When  approached 
from  the  side  they  are  much  more  easily  alarmed  and  fly  away.  When  approached  from 
above,  they  do  not  move  until  the  stick,  finger,  or  whatnot,  is  within  a  couple  of  inches, 
when  they  remove  the  fore  feet  from  the  support  and,  on  nearer  approach,  fall  to  the  ground. 
When  in  copulation,  the  female  tries  to  disengage  by  rapidly  vibrating  the  wings  in  attempted 
flight,  repeating  this  often,  from  every  one  to  five  seconds  until  disengaged  or  exhausted. 
The  male  can  disconnect  himself  at  will.  In  copulation  the  female  is  always  uppermost 
unless  exhausted,  when  both  sexes  lie  flat  on  a  grass-blade.  The  female  has  the  head  up, 
the  male  the  head  downward;  copulation  always  takes  place  on  a  vertical  support,  usually 
a  blade  of  grass,  sometimes  a  plant  stem.  The  sexes  remain  in  copulation  for  quite  a  long 
time  and  are  perfectly  motionless.  All  of  the  legs  of  both  sexes  are  on  the  support  unless 
in  a  position  where  this  is  physically  impossible,  in  which  case  as  many  as  possible  are  used; 
the  hind  legs  of  both  sexes  are  held  at  right  angles  to  the  support,  the  forelegs  in  front.  After 
copulation  the  female  generally  drops  to  the  ground,  the  male,  after  a  few  moments'  rest,  flies 
away.  Specimens  in  copulation  were  found  in  abundance  from  2  to  4.30  p.m.  when  no  more 
could  be  discovered.  From  4.30  to  7  p.m.  solitary  males  were  common,  but  no  females  could 
be  found  on  the  grass-blades.  At  4.30  p.m.,  a  few  females  were  found  clinging  to  the  trunks 
of  the  willow  trees  about  two  feet  from  the  ground.  At  5.30  p.m.,  females  were  noted  in 
small  groups  over  the  water,  evidently  engaged  in  oviposition,  as  they  frequently  dipped 
down  to  the  surface.  These  latter  were  in  company  with  a  large  swarm  of  dancing  empidid 
flies  (Rhamphomyia).  Of  the  great  numbers  that  were  picked  from  grasses  in  the  after- 
noon a  considerable  proportion  were  females  and  toward  5  p.m.  they  commenced  egg-laying 
on  the  sides  of  the  containing  vessel,  large  shell  vials.  By  7  p.m.  the  sides  of  the  vials  were 
black  in  places  with  the  large,  dark-colored  eggs.  This  data  would  seem  to  place  the  time 
for  oviposition  at  about  sunset.  The  females  are  very  good  fliers  and  often  travel  for  long 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  879 

distances  before  alighting  as  is  shown  toward  sunset  when  they  fly  for  long  stretches  up- 
stream. The  males  are  rather  poor  fliers,  due  in  part,  possibly,  to  the  weight  of  the  long 
antennae  and,  as  stated  before,  this  sex  prefers  to  skulk  rather  than  fly.  When  the  males 
fly,  they  do  so  heavily  and  seize  the  first  support  that  they  collide  with  and  hang  on,  occasion- 
ally flying  on  immediately  to  another  support.  When  the  male  comes  in  contact  with  a  stem, 
he  very  often  ascends  to  the  top  by  means  of  a  part-flying,  part-climbing  motion  and,  on  reach- 
ing the  summit,  flies  off  to  another  place.  As  a  rule  the  flies,  especially  the  females,  alight  on 
a  single  grassblade,  but  very  often  the  males  are  observed  on  two  blades,  the  legs  of  one  side 
on  one  blade  and  those  of  the  opposite  side  on  the  other;  when  the  body  thus  hangs  between 
the  stalks,  the  tarsi  diverge  from  one  another,  whereas  on  a  single  support,  the  legs  converge. 

Hexatoma  was  preyed  upon  by  large  numbers  of  a  scatophagid  fly  that  occurred  in  great 
abundance  in  this  vicinity  and  seemed  to  be  subsisting  almost  entirely  on  these  (flies.  At 
least  twenty  of  these  predaceous  flies  were  noted  with  Hexatomae  and  this  species  seemed 
to  constitute  the  principle  insect  enemy  of  the  crane-fly;  'They  would  lurk  on  the  grass 
blades  and  sally  forth  after  their  prey,  carrying  it  back  to  some  point  to  feed  upon  it.  It  is 
probable  that  the  blood  is  taken  since  the  body  of  the  Hexatoma  appeared  almost  uninjured 
when  examined.  On  an  old  beam  where  males  had  a  habit  of  walking  up  the  vertical  face, 
a  small  spider's  web  was  found,  in  which  eleven  specimens  were  entangled,  two  being  still 
alive;  eight  of  these  were  males,  the  remaining  three,  females. 

The  males  especially  can  walk  up  smooth  surfaces,  as  glass,  moving  the  legs  alternately 
and  awkwardly.  The  first  pair  taken  were  in  copulation  but  in  placing  them  in  the  vial 
they  became  disengaged  and  ran  about  in  the  container.  After  a  short  time  they  began  to 
copulate  in  the  tube. 

May  15,  1911  — A  pair  were  taken  in  copulation  at  10  a.m.;  at  8  p.m.  they  were  still  in 
coitu,  but  this  is  exceptional  as  most  of  the  pairs  disengage  very  readily.  In  the  morning 
the  species  is  very  active  and  although  the  males  do  not  fly  far,  they  fly  readily  and  it  is 
difficult  to  pick  them  up  by  hand.  The  females  are  excellent  fliers  especially  in  the  morning. 

Several  pairs  were  taken  in  copulation  and  each  pair  was  isolated  in  a  separate  vial  in  order 
to  ascertain  the  number  of  eggs  per  female.  The  clutch  was  determined  by  dissection. 
When  the  captive  insects  began  to  oviposit,  the  eggs  shot  out  from  the  body,  at  first  slowly, 
then  more  rapidly,  one  per  second,  later  much  slower  again,  the  eggs  being  extruded  one  at 
a  time.  The  total  period  of  ovi position  required  seven  minutes;  at  the  end  of  sixty  seconds, 
in  the  space  between  sixty  and  seventy  seconds,  eighteen  eggs  were  laid,  or  1.8  per  second. 
Toward  the  end  of  egg-laying,  the  eggs  appeared  much  more  slowly,  one  in  two  seconds.  The 
eggs  are  quite  sticky  or  viscid  and  adhere  to  the  glass.  When  the  female  is  in  -danger  of 
death,  as  when  she  falls  into  the  water,  she  begins,  at  once,  to  deposit  the  egg-complement. 
In  nature  it  seems  probable  that  one  egg  is  laid  at  each  descent  to  the  water.  As  soon  as 
the  female  touches  the  water,  although  she  has  not  deposited  an  egg  all  day,  she  immediately 
starts  to  deposit  the  oblong  black  eggs.  After  the  last  egg  is  expelled  the  muscles  of  the 
ovipositor  still  go  through  the  motions  of  expulsion.  One  specimen  was  placed  in  the  water 
and  as  usual  began  to  deposit  her  eggs.  She  was  decapitated,  laid  eleven  eggs  and  tried  to 
lay  still  more  but  failed.  The  number  of  eggs  laid  varied  from  316  to  372  with  an  average 
of  347;  the  time  required  for  oviposition  varied  from  seven  minutes  to  seven  minutes  and  forty 
seconds.  In  most  cases  the  number  of  the  egg-complement  is  probably  between  300  and  400. 

•  "  <4 

The  greater  part  of  the  larval  existence  is  probably  spent  in  the  water, 
and  it  is  only  when  the  larvae  are  fully  grown  and  ready  to  pupate  that 
they  come  to  land.  On  April  26,  1914,  gravel  from  the  bank  of  Casca- 
dilla  Creek  was  carefully  examined,  but  no  signs  of  larvae  or  pupae  were 
to  be  discovered.  On  May  6,  however,  the  same  bank  was  examined 
and  about  ten  larvae  and  seventy-five  pupae  were  found.  Sometimes  the 
immature  stages  are  very  abundant.  On  May  12,  1917,  near  the  place 


880  CHAELES  PAUL  ALEXANDER 

just  described,  the  writer  found  larvae  and  newly  transformed  pupae  in 
great  numbers,  the  former  pupating  in  the  dry  sand  rather  distant  from 
the  water's  edge.  In  one  dry  patch  of  sand  on  a  rocky  ledge,  sixty 
specimens  were  found  in  six  square  inches  of  soil.  The  insects  are  often 
found  in  gravel  or  coarse  sand  that  is  thickly  penetrated  by  grass  roots 
and  rhizomes,  rarely  in  pure  gravel.  They  are  most  commonly  found 
in  soil  that  has  been  recently  deposited  after  freshets.  The  immature 
stages  of  Hexatoma  are  associated  with  larvae  of  Eriocera  spinosa,  E. 
dnerea,  Tipula  bella,  and  Atherix  probably  variegata  Walk.,  with  pupae 
of  Chrysops  excitans  Walk.,  and  with  many  beetles  such  as  Paederus 
littorarius  Grav.,  Gastrolobium  bicolor  (Grav.),  Bledius  sp.,  Omophron  sp., 
Dyschirius  sphaericollis  Say,  Tachistodes  partiarius  (Say),  Anadaptus 
discoideus  (Dej.),  Laccobius  agilis  Rand,  and  other  species  characteristic 
of  the  sandy  margins  of  large  streams.  The  pupal  duration  of  Hexatoma 
is  six  and  one-half  days. 

Larva. —  Length,  14-15  mm. 
Diameter,  1-1.3  mm. 

Color  of  body,  pale  brownish  yellow. 

Body  covered  with  rather  abundant  appressed  hairs,  the  subterminal  distended  part  of 
abdomen  with  transverse  rows  of  very  short  spines  or  setae,  there  being  from  thirty-five  to 
forty  such  rows.  Spiracular  disk  (Plate  LI,  246)  surrounded  by  four  lobes;  ventral  lobes 
the  longer,  with  a  fringe  of  a  few  long  hairs  at  apex;  on  inner  face  a  long,  narrow,  brown 
mark,  extending  from  the  tip  inward,  the  proximal  end  expanded;  lateral  lobes  with  numerous 
long  hairs  which  are  gradually  shorter  toward  base  of  lobes.  Spiracles  circular,  situated  at 
base  of  lateral  lobes;  a  dusky  mark  extending  from  spiracles  dorsad. 

Head  capsule  of  usual  hexatomine  type,  as  described  for  Eriocera  longicornis  (page  888). 
Entire  general  features  and  details  of  mouth  parts  very  similar  to  those  of  Eriocera  longicornis. 
Labrum  (Plate  LI,  243)  transversely  oval,  with  frame  strong  and  chitinized,  anterior  median 
part  produced  into  a  small  lobe  (Plate  LII,  250)  bearing  two  lateral  papillae  and  two  setiferous 
tubercles  surrounded  by  short  hairs;  lateral  margins  of  labrum  produced  into  prominent 
lobes  directed  proximad  and  cephalad,  densely  hairy  and  entirely  protecting  anterior  margin 
of  labrum.  Antenna  (Plate  LI,  244)  with  apical  papillae  shorter  than  segment,  the  largest 
papilla  transversely  sculptured  (Plate  LII,  251).  Mandible  (Plate  LI,  245)  long  and  slen- 
der, the  lateral  teeth  more  accentuated  than  in  Eriocera;  largest  tooth  with  a  flattened  trun- 
cated blade  in  its  axil,  this  in  some  cases  broken  up  into  two  or  three  small  blades;  basad 
of  largest  lateral  tooth  a  flattened  lobe  which  is  barely  indicated  in  the  species  of  Eriocera 
studied.  Maxilla  with  outer  flattened  blade  conspicuous;  palpus  near  its  base  on  inner 
side,  similar  to  the  condition  obtaining  in  Eriocera;  just  laterad  of  palpus  a  powerful  seta. 

Pupa.—  Length,  9.2-9.6  mm. 

Width,  d.-s.,  1.2  mm.     v 
Depth,  d.-v.,  1.4  mm.    ] 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  881 

Pupa  (Plate  LI,  247)  very  similar  to  that  of  Eriocera  longicornis,  differing  only  in  its  small 
size,  greater  development  of  scapal  spine,  lack  of  projection  on  mesonotal  prescutum,  and 
a  few  lesser  characters.  Cephalic  crest  (Plate  LI,  248  and  249)  as  viewed  from  beneath, 
very  different  in  shape.  Fore  pair  of  legs  much  shorter  than  the  others,  ending  just  beyond 
posterior  margin  of  second  abdominal  segment;  hind  pair  of  legs  extending  far  beyond  the 
others,  ending  beyond  midlength  of  third  abdominal  segment;  in  some  specimens  the  tarsal 
segments  much  closer  to  posterior  margin  of  third  abdominal  segment,  but  usually  a  marked 
difference  in  tips  of  tarsi  of  the  various  legs.  Cephalic  crest  as  viewed  from  side,  triangular, 
ending  in  an  acute  point  directed  strongly  forward.  Viewed  from  beneath,  lobes  conspicu- 
ously triangular,  pointed,  lying  parallel  or  slightly  divergent  and  separated  by  a  deep 
median  split.  Spine  on  scape  of  antennae  very  large,  conspicuous.  Tubercle  on  labrum 
strongly  developed.  Antennal  sheaths  of  male  very  long,  those  of  female  much  shorter. 
Pronotal  breathing  horns  short,  straight.  Mesonotum  strongly  wrinkled  along  median 
line;  scutellar  lobe  not  developed.  Wing  pads  dark,  venation  not  showing  clearly  but,  if 
made  out,  the  very  short  cell  /fo  and  the  reduced  M  characteristic  of  Hexatoma  alone. 
Posterior  leg  sheaths  extending  beyond  level  of  middle  legs,  these,  in  turn,  being  longer 
than  sheaths  of  fore  legs. 

Abdomen  (Plate  Lit,  252)  with  about  thirty-four  spicules  on  sternite  4,  about  thirty 
in  a  straight,  uninterrupted  row  on  tergites  3  and  4,  and  about  twenty  on  tergite  5. 
Chaetotaxy  about  as  in  Eriocera  longicornis,  but  the  seta  lying  ventrad  of  spiracle  on  pleurites 
much  farther  ventrad  and  very  weak.  Male  cauda  with  sternal  lobes  strongly  rounded, 
enlarged,  and  bent  suddenly  dorsad.  Female  cauda  as  that  of  male;  ovipositor  viewed 
from  side  with  an  obtuse  notch;  from  beneath,  sternite  obtusely  pointed  and  with  a  deep 
median  split;  from  above,  tergite  almost  flat  across  caudal  margin,  the  lateral  angles  rounded, 
with  a  deep  median  split. 

Nepionotype. —  Ithaca,  New  York,  May  6,  1914. 

N&anotyve. —  Type  locality,  May  2,  1913. 

Paratyjms. —  Several  hundred  larvae  and  pupae  from  type  locality. 

Genus  Eriocera  Macquart  (Gr.  wool  +  horn) 

1830  Caloptera  Guer.     Voyage  de  la  Coquille,  Zool.,  Ins.,  pi.  20,  fig.  2. 

1838  Eriocera  Macq.     Dipt.  Exot.,  vol.  1,  p.  74. 

1838  Evanioptera  Gue>.     Voyage  de  la  Coquille,  Zool.,  vol.  2,  part  2,  p.  287. 

1848  Pterocosmus  Walk.     List  Dipt.  Brit.  Mus.,  vol.  1,  p.  78. 

1850  Allarithmia  Loew.     Bernstein  und  Bernsteinfauna,  p.  38. 

1857  Oligomera  Dolesch.     Natuurk.  Tijdschr.  Nederl.  Indie,  vol.  14,  p.  387. 

1859  Arrhenica  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  242. 

1859  Physecrania  Bigot.     Ann.  Soc.  Ent.  France,  ser.  3,  vol.  7,  p.  123,  pi.  3,  fig.  1. 

1912  Androclosma  Enderlein.     Zool.  Jahrb.,  vol.  32,  part  1,  p.  34. 

1916  Globericera  Matsumura.     Thous.  Ins.  Japan,  add.  2,  p.  471. 

Larva.—  Spiracular  disk  surrounded  by  four  lobes  which  are  rarely  (as  in  Eriocera  cinerea, 
subobsolete,  inner  face  lined  with  brown  or  black,  tips  with  fringes  of  moderately  long  hairs. 
Head  capsule  long,  narrow,  the  constituent  plates  very  slender;  dorsal  plate  completely 
divided  tho  contiguous  or  approximated  behind.  Labral  sclerite  large  and  conspicuous, 
sensory  tubercles  and  papillae  crowded  on  median  cephalic  region.  Mandible  long,  acute 

7 


882  CHARLES  PAUL  ALEXANDER 

at  tip,  with  two  teeth  at  about  midlength.  Maxilla  with  outer  lobe  greatly  prolonged  into 
a  flattened  blade.  Antenna  cylindrical  or  clavate,  with  three  Or  four  long  papillae  at  tip. 
Mentum  not  chitinized,  in  E.  cinerea  with  a  flattened  rectangular  plate  on  either  side,  this 
armed  with  numerous  hooks  and  spines. 

Pupa.- — Cephalic  crest  of  various  shapes  and  sizes,  very  reduced  in  Eriocera  spinosa. 
Antennal  sheaths  of  males  of  several  species  (E.  spinosa,  E.  longicornis,  E.  cinerea)  very 
long,  extending  beyond  end  of  wing  pad.  Pronotal  breathing  horns  of  various  shapes,  acutely 
pointed  in  E.  spinosa,  short  and  blunt  in  several  species.  Head  and  thorax  often  wit'i 
spines  or  tubercles  on  scape  of  antenna,  on  labrum,  or  (in  E.  spinosa)  on  face  of  eye;  a 
tubercle  on  scutellum  (in  E.  longicornis),  one  on  dorsum  of  second  abdominal  segment  (in  E. 
spinosa).  Abdominal  segments  with  a  conspicuous  transverse  armature  of  spines  near 
posterior  margin.  Lateral  spiracles  large,  distinct. 

Eriocera  is  an  extensive  genus  (including  approximately  150  species) 
of  medium-sized  to  large-  flies,  most  of  which  are  tropical.  The  genus 
has  not  been  found  in  Europe,  but  elsewhere  it  is  represented  by  a  host 
of  species.  The  habits  of  the  adult  flies  have  already  been  noted  (page 
704).  The  immature  stages  are  spent  in  sand  or  gravel  near  running 
water,  more  especially  along  large  streams.  A  more  complete  account 
of  the  genus  is  given  by  Alexander  and  Lloyd  (1914)  and  by  Alexander 
(1915  c:  148-152). 

The  occurrence  of  the  flies  is  somewhat  local.  During  an  entire  summer 
of  collecting  in  Maine  in  1913,  the  writer  did  not  find  a  single  specimen 
of  any  species;  and  Dr.  Dietz  has  stated  that  the  only  living  individual 
which  he  has  found  was  a  single  male  of  Eriocera  spinosa  takeji  in  the 
Pocono  Mountains  in  Pennsylvania.  On  the  other  hand,  the  flies  are 
often  found  in  countless  numbers,  and  several  species  may  be  found 
associated  together.  Thus,  at  Ithaca,  New  York,  in  the  sandy  gravel 
along  Cascadilla  Creek,  the  immature  stages  of  four  species  of  Eriocera 
and  one  of  the  closely  related  Hexatoma  occur  together  in  unlimited 
numbers. 

The  species  of  Eriocera  may  be  separated  by  the  following  keys: 

Larvae 

1.  Lobes  surrounding  spiracular  disk  obsolete  or  nearly  so;  a  flat,  chitinized  plate  with  ser- 

rate margins  on  either  side  of  mental  region E.  cinerea  Alex.  (p.  886) 

Spiracular  disk  surrounded  by  four  slender  lobes;  no  plate  as  described  above  on  mental 
region 2 

2.  Very  large  (length  40-45  mm.,  diameter  4-5  mm.);  spiracular  disk  with  ventral  lobes 

narrowly  lined  with  black,  inner  ends  of  each  forked,  Y-shaped;  lateral  lobes  narrowly 

\    lined  with  black,  inner  ends  of  marks  expanded E.  spinosa  (O.  S.)  (p.  883) 

Smaller  (length  under  30  mm.,  diameter  under  2.5  mm.) ;  spiracular  disk  not  marked 
as  above 3 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  883 

3.  Ventral  lobes  of  spiracular  disk  bearing  one  or  two  very  long,  dark  setae  in  addition  to 
the  shorter  yellowish  fringe;  inner  face  of  each  lobe  with  a  capillary  black  line  which 
is  suddenly  expanded  at  its  inner  end  into  a  triangular  brown  mark,  the  two  marks 
inclosing  an  oval  pale  area  between  their  proximal  ends;  lateral  lobes  with  a  capillary 

black  line E.  fultonensis  Alex.  (p.  890) 

Ventral  lobes  of  spiracular  disk  with  apical  fringe  consisting  of  numerous  long,  pale  setae; 
inner  face  of  each  lobe  lined  with  pale  brown,  at  about  midlength  gradually  expanded 
into  an  elongate-triangular  mark,  the  two  marks  inclosing  a  linear  pale  area  between 
their  proximal  ends;  lateral  lobes  with  a  brown  line E.  longicornis  (Walk.)  (p.  888) 

Pupae 

1.  Size  large  (length  25  mm.  or  over);  pronotal  breathing  horns  tapering  to  acute  tips; 

cephalic  crest  small,  reduced  to  four  small  tubercles;  cell  MI  on  wing  pad  present; 
a  strong  spinous  tubercle  on  either  side  of  median  line  at  base  of  second  abdominal 

tergite;  a  tubercle  on  eye E.  spinosa  (0.  S.)  (p.  883) 

Size  smaller  (length  under  18  mm.);  pronotal  breathing  horns  blunt  at  tips;  cephalic 
crest  prominent;  cell  MI  on  wing  pad  lacking;  no  tubercles  on  second  abdominal  tergite 
or  on  eye 2 

2.  A  tubercle  on  mesonotal  scutellum E.  longicornis  (Walk.)  (p.  888) 

No  tubercle  on  mesonotal  scutellum 3 

3.  Pleurites  of  abdominal  segments  with  a  transverse  row  of  three  setae  ventrad  and  slightly 

caudad  of  spiracle;  antennae  of  male  elongated E.  cinerea  Alex.  (p.  886) 

Pleurites  of  abdominal  segments  with  two  stout  setae  dorsad  and  caudad  of  spirarh; 
antennae  short  hi  both  sexes E.  fultonensis  Alex.  (p.  890) 

Eriocera  spinosa  (0.  S.) 

1859     Arrhenica  spinosa  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  244. 

Eriocera  spinosa  O.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  252-253. 


Eriocera  spinosa  is  the  commonest  of  the  large  species  of  the  genus 
in  eastern  North  America.  The  larvae  occur  in  great  numbers  beneath 
rocks  in  rapid  water  in  the  autumn,  when  they  form  a  considerable  pro- 
portion of  the  insect  life  in  the  streams.  When  about  to  pupate  they  go 
to  the  neighboring  banks  and  live  for  some  time  in  the  sand  or  gravel. 
The  habits  of  the  larvae  have  been  discussed  by  Alexander  and  Lloyd 
(1914:16-17)  and  by  Alexander  (1915c:149). 

The  larvae  were  found  on  May  1,  1913,  along  the  banks  of  Fall  Creek, 
Ithaca,  New  York,  in  considerable  numbers.  They  were  associated  with 
young  and  mature  pupae  of  E.  longicornis,  which  were  emerging  in  great 
numbers  at  the  time.  On  May  27,  both  larvae  and  pupae  of  E.  spinosa 
were  found  to  be  very  abundant,  the  larvae  being  more  numerous  in  the 
wetter  places,  the  pupae  in  the  drier  spots.  They  occurred  at  various 
distances  from  the  water's  edge,  from  within  a  foot  of  the  margin  to  as 
far  back  as  eight  or  ten  feet.  The  pupae  are  found  in  short,  more  or  less 
vertical,  burrows,  from  one  to  three  inches  below  the  surface.  Not  often 


884  CHARLES  PAUL  ALEXANDER 

were  larvae  and  pupae  found  in  close  proximity  to  each  other.  Pupae 
of  E.  spinosa,  as  well  as  of  all  other  species  of  the  tribe  as  known,  are 
very  active  when  removed  from  their  burrows,  wriggling  rapidly  to  and 
fro,  and  are  exceedingly  tenacious  of  life.  Larvae,  as  found  on  May 
27,  were  mostly  contracted;  a  few,  however,  were  expanded  and  had  the 
subterminal  segment  of  the  abdomen  swollen.  In  this  regard  it  may  be 
mentioned  that  almost  all  of  the  larvae  of  crane-flies  that  live  in  the  sand 
or  mud  along  the  banks  of  streams  have  this  ability  to  inflate  the  end 
of  the  abdomen.  Larvae  of  Eriopterini,  of  Pediciini,  and  of  Hexatomini 
have  been  observed  with  this  conspicuous  enlargement.  It  is  undoubtedly 
used  to  propel  the  larva  thru  the  soil  by  alternate  expansion  and  con- 
traction of  the  segment. 

Larvae  of  E.  spinosa  were  placed  in  breeding  jars  on  May  13  and  adult 
flies  emerged  on  the  28th.  It  is  probable  that  the  pupal  stage  is  not 
longer  than  from  ten  to  twelve  days,  at  the  most.  On  May  30  a  large 
number  of  larvae  and  pupae  were  brought  into  the  laboratory  in  a  bucket 
of  gravel.  Some  of  the  fully  matured  pupae  transformed  in  the  pail 
while  being  brought  to  the  laboratory. 

The  larvae  are  carnivorous.  Their  powerful,  sickle-shaped  mandibles 
are  capable  of  inflicting  a  painful  bite  on  tender  parts  of  the  hand.  S.  G. 
Rich  placed  larvae  in  dishes  together  with  the  nymphs  of  various  dragon 
flies.  The  smaller  of  the  nymphs  were  eaten  by  the  Eriocera  larvae,  thus 
confirming  previous  observations  on  the  carnivorous  habits  of  the  species. 

Larva. —  Length  when  fully  extended,  40-45  mm. 
Diameter,  4-5  mm. 

Color  varying  from  very  pale  whitish  to  rather  dark  brown;  in  life,  the  skin  showing 
conspicuous  bronzy  reflections. 

Spiracular  disk  (Plate  LI V,  262)  surrounded  by  four  slender  elongate  lobes,  one  pair  being 
lateral,  the  other  ventral,  in  position;  inner  face  of  lateral  lobes  with  a  capillary  black  line, 
this  beginning  as  an  enlarged  black  spot  just  ventrad  of  spiracle,  reaching  tip  of  lobe;  dorsal 
outer  edge  of  lobe  with  a  dense  fringe  of  long,  conspicuous,  reddish  hairs,  inner  edge  of  row 
beginning  just  laterad  of  spiracle  where  the  hairs  are  very  short,  gradually  becoming  longer 
to  tip,  where  they  are  as  long  as  the  lobe  itself;  ventral  lobe  with  a  capillary  black  line  on 
proximal  edge,  this  dividing  at  base  of  lobe,  the  lower  branch  running  along  ventral  margin 
of  stigmal  field  and  approaching  its  fellow  of  the  opposite  side  on  median  line  of  body;  a 
dense  fringe  of  conspicuous  reddish  hairs  at  tip  of  lobe  and  continued  on  outer  dorsal  side 
for  a  short  distance  toward  base;  a  few  dusky  brown  spots  on  stigmal  field  between  spiracles; 
two  small  hairs  between  spiracles.  Spiracles  rather  small,  widely  separated.  Underneath 
caudal  lobes  and  behind  penultimate  swollen  segment,  four  anal  gills,  short,  stout,  cylindrical, 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  885 

the  lateral  pair  directed  outward,  the  inner  pair  directed  caudad.  Head  capsule  (Plate 
LIV,  267)  broad  in  proportion  to  its  length,  measuring  3.5-3.8  mm.  by  1.8-2  mm.  (across 
dorsal  plates).  Papillae  at  tip  of  antenna  short,  not  more  than  one  third  length  of  seg- 
ment. Mandible  (Plate  LIV,  271)  lacking  a  prominent  conical  tooth  at  midlength,  such  as 
is  found  in  E.  cinerea  and  other  species.  1  < 

Pupa. —  Length:  male,  26.5-27  mm.;  female,  25-28.5  mm. 

Width,  d.-s.:  male,  3.4-3.9  mm.;  female,  3.4-4  mm. 
Depth,  d.-v.:  male,  4-4.2  mm.;  female,  3.5-4  mm. 

In  life,  pupae  varying  in  color  from  very  pale  yellowish  to  dark  brown  or  almost  black, 
the  deepest  color  being  that  of  head  and  thorax  of  old  pupae;  body  often  showing  bronzy 
reflections. 

Cephalic  crest  very  reduced,  scarcely  projecting  beyond  level  of  antennae;  viewed  from 
beneath,  somewhat  quadrate,  the  anterior  lateral  angles  produced  into  small  pointed  lobes 
bearing;  a  small  seta  at  apex;  viewed  from  side  a  second  pair  of  lobes  is  seen,  these  being 
subequal  to  anterior  lobes  in  size,  and  likewise  setiferous.  Spine  of  antennal  scape  very 
large,  somewhat  curved,  directed  ventrad.  Inner  caudal  surface  of  eye  with  a  conspicuous 
tubercle.  Tentorial  region  produced  into  a  small  median  tubercle.  Tubercles  at  base  of 
labrum  very  large,  close  together,  their  tips  strongly  chitinized;  a  small  seta  above  each 
of  these  labral  tubercles  and  another  small  seta  on  each  cheek.  Pronotal  breathing  horns 
long,  slender,  broad  at  base,  flattened  and  rather  pointed  at  tips,  the  organ  arcuated  so  that 
apex  is  bent  strongly  ventrad.  Mesonotal  scutellar  lobe  (Plate  LV,  280)  prominent,  rather 
strongly  projecting.  Wing  pad  light  brown,  venation  showing  very  clearly,  the  presence  of 
cell  MI  in  connection  with  elongate  antennae  in  male  sex  being  found  in  this  species  alone 
in  eastern  North  America.  Leg  sheaths  with  tarsal  sheaths  ending  on  a  level,  about  opposite 
end  of  third  abdominal  segment. 

Second  abdominal  tergite  with  a  conspicuous  basal  tubercle  on  either  side  of  median  line. 
Abdominal  segments  (Plate  LVI,  283)  with  subterminal  rows,  of  spines,  there  being  about 
twenty  to  twenty-two  on  tergites  2  to  5;  tergites  6  and  7  destitute  of  spines  but  with  four 
subapical  setiferous  tubercles;  tergites  2  to  7  with  a  conspicuous  setiferous  tubercle  on  ventro- 
cephalic  angle  of  each  posterior  ring;  eighth  tergite  concave  on  posterior  margin,  bearing  a 
pair  of  strong  apical  tubercles  on  either  side  of  median  line.  Pleural  region  of  abdomen 
rather  restricted,  longitudinally  wrinkled.  Spiracles  large,  elliptical,  transverse,  placed  about 
opposite  midlength  of  segments.  Three  small  setiferous  tubercles  ventrad  and  caudad  of 
spiracle,  and  another  similar  tubercle  on  dorsa-cephalic  angle  of  each  pleuron.  Semites 
on  segment  3  with  two  spines  on  each  outer  angle;  segments  4  to  6  with  from  sixteen  to  twenty 
spines;  segment  7  with  about  ten  spines;  an  isolated  set'ferous  tubercle  caudad  and  dorsad 
of  ends  of  row;  segments  4  to  7  with  a  setiferous  tubercle  about  midlength  of  posterior  ring; 
segment  8  lacking  soft  pleural  region,  bearing  an  apical  row  of  strong  spines  which  are  inter- 
rupted only  on  dorsum  and  for  a  small  space  on  median  line  of  venter,  there  being  about 
twenty  of  these  spines  in  the  circlet.  Male  cauda  (Plate  LV,  281  and  282)  with  ninth  sternite 
rounded,  swollen,  with  a  deep  median  furrow  bearing  a  small  lobe  on  ventral  side  at  end  of 
split;  ninth  tergite  produced  caudad  into  two  strong  conical  points  separated  by  a  V-shaped 
notch,  these  points  directed  caudad  and  slightly  dorsad,  each  one  a  little  split  near  tip  on 
outer  face  and  with  a  prominent  lateral  tooth  at  about  midlength.  Female  cauda  (Plate 


886  CHARLES  PAUL  ALEXANDER 

LVI,  284)  with  ninth  sternite  elongated,  cylindrical,  its  tip  rounded,  feebly  split  beneath; 
ninth  tergite  very  long,  pointed,  with  a  deep  median  split. 

Nepionotype. —  Ithaca,  New  York,  May  1,  1913. 

Neanotype. —  With  type  larva. 

Paratypes. —  Numerous  larvae  and  pupae  witji  types,  May  1-15,  1913. 

Eriocera  cinerea  Alex. 

1912    Eriocera  cinerea  Alex.    Psyche,  vol.  19,  p.  169-170,  pi.  13,  fig.  9. 

Eriocera  cinerea  is  locally  common,  flying  in  May.  The  larvae  were 
found  on  May  16,  1917,  in  sand  along  the  banks  of  Cascadilla  Creek, 
Ithaca,  New  York.  They  have  been  found  at  various  dates  during  the 
past  few  years,  but  always  in  scanty  numbers.  Larvae  found  on  April 
28  were  associated  with  larvae  of  Hexatoma  megacera,  Eriocera  spinosa, 
Atherix,  and  other  insects.  The  larvae  are  stouter  than  those  of  E, 
longicornis  and  are  pale  whitish  yellow,  quite  devoid  of  the  greenish  tints 
of  the  latter  species.  A  larva  found  on  April  28  transformed  to  an  adult 
female  on  May  16.  Additional  larvae  and  pupae  were  found  on  May 
24,  and  a  few  pupae  on  May  30,  1917. 

The  supposed  larva  of  E.  longicornis  described  by  Alexander  and  Lloyd 
(1914:21-23)  pertains  to  this  species;  the  true  longicornis  is  discussed 
later  in  this  paper. 

Larva. —  Length,  15-16  mm. 
Diameter,  2-2.2  mm. 

Color  light  yellow. 

Form  almost  terete,  abdominal  segments  subdivided  into  two  annuli.  Subterminal 
abdominal  segment  greatly  enlarged,  capable  of  great  distention.  Spiracular  disk  (Plate 
LIV,  263  and  264)  very  reduced,  the  usual  four  lobes  exceedingly  small;  ventral  lobes  prac- 
tically obsolete,  not  projecting,  each  fringed  with  from  twenty  to  thirty  long,  golden-yellow 
hairs;  lateral  lobes  very  short,  triangular,  fringed  with  from  twelve  to  fifteen  long  hairs; 
a  faint  dusky  mark  from  dorsal  margin  of  each  spiracle  to  edge  of  field;  a  faint  vertical  stripe 
between  spiracles;  ventral  lobes  marked  with  brownish  black,  the  mark  of  each  side  three- 
pointed  at  its  inner  end,  the  innermost  of  these  points  connected  with  its  fellow  of  the 
opposite  side;  lateral  lobes  with  the  marks  elongate,  triangular,  the  points  directed  outward. 
Spiracles  small,  oval,  separated  by  a  distance  a  little  less  than  the  diameter  of  one.  Anal 
gills  four,  very  short  and  inconspicuous. 

Head  capsule  long  and  narrow,  measuring  about  1.5  by  0.275  mm.;  dorsal  plates  of  capsule 
with  proximal  anterior  angles  produced  inward.  Labral  sclerite  (Plate  LII,  253)  having 
labrum  itself  subquadrate.  Mental  region  entirely  lacking  strongly  chitinized  points  as  in 
this  group  of  genera.  Present  species  showing  a  structure  which  is  probably  a  part  of  labium, 
either  mentum  or  hypopharynx,  and  which  has  not  been  found  in  any  other  species  of  the 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  887 

genus  (Plate  LII,  254) ;  this'structure  located  on  either  side  of  capsule  on  ventral  face,  a 
flattened,  subrectangular  plate  whose  surface,  except  at  base,  is  densely  set  with  small  spines 
and  large  pits;  inner  margin  provided  with  large,  acute  spines,  beginning  at  about  one- 
third  length  of  sclerite,  gradually  enlarged  toward  tip,  at  inner  angle  acute;  these  teeth  inter- 
rupted before  outer  posterior  angle,  which  terminates  in  a  blunt,  flattened  lobe.  Antenna 
cylindrical,  apex  obliquely  truncated,  terminating  in  a  slender  apical  papilla  which  is  longer 
than  the  segment  that  bears  it,  broad  at  base,  tapering  gradually  to  tip;  two  or  three  long 
apical  setae;  small  auditory  plates  at  about  one-third  length  of  segment.  Mandible  (Plate 
LII,  255)  a  powerful,  slender,  curved  hook,  at  about  midlength  with  a  strong  pointed  tooth 
bearing  in  its  axil  a  smaller  tooth;  as  is  usual  in  the  genus,  an  egg-shaped  chitinized  piece 
isolated  in  one  of  the  ventral  tendons  of  mandible.  Maxilla  arising  just  ventrad  of  man 
dible,  outer  lobe  persisting  as  a  very  elongate,  bladelike  organ. 

Pupa. —  Length,  13-15  mm. 
Width,  d.-s.,  1.6  mm. 
Depth,  d.-v.,  1.7  mm. 

Head  and  appendages  dark  brown,  thoracic  dorsum  a  little  paler;  abdomen  with  posterior 
rings  of  tergum  and  sternum  dark  brown,  producing  a  banded  appearance. 

Cephalic  crest  (Plate  LIII,  257)  very  large  and  conspicuous,  consisting  of  two  rounded 
lobes  behind,  each  tipped  with  a  strong  seta;  anterior  part  of  crest  directed  ventrad  and  con- 
sisting of  two  lobes,  the  larger  bearing  a  strong  seta  on  outer  face;  viewed  from  front,  these 
anterior  lobes  separated  by  a  very  narrow,  U-shaped,  median  notch;,  crest  of  female  a  little 
smaller.  Two  blunt  tubercles  on  scapal  segments  of  each  antenna.  Junction  of  clypeus  and 
labrum  with  two  tubercles,  above  and  slightly  laterad  of  each  a  strong  seta.  A  strong  seta 
on  cheek  below  eye.  Labrum  broad,  very  obtusely  rounded  at  tip.  Labial  lobes  rectangu- 
lar, widely  separated.  Maxillary  palpi  blunt  at  tips  (Plate  LIII,  258).  Antenna  of  male  elon- 
gate, exceeding  wing  and  ending  opposite  base  of  last  tarsal  segment  of  hind  leg.  Pronotal 
breathing  horns  short,  slender,  cylindrical,  scarcely  longer  than  cephalic  crest.  Two  long 
curved  setae  and  a  smaller  straight  seta  laterad  and  ventrad.  of  base  of  each  breathing  horn. 
Lateral  angle  of  thorax  with  two  setae;  two  long  setae  above  wing  axil  (Plate  LIII,  256). 
A  strong  seta  on  either  side  of  mesonotum  and  a  group  of  two  small  setae  in  front  of  each 
of  these.  Wing  sheaths  extending  to  base  of  third  abdominal  segment.  Leg  sheaths  extend- 
ing to  base  of  fourth  abdominal  segment;  tarsal  sheaths  ending  about  on  a  level,  or  those 
of  fore  legs  considerably  shorter. 

Abdominal  segments  (Plate  LIII,  259)  divided  into  two  subequal  rings.     Chaetotaxy  as 
follows:  pleura  with  a  seta  on  dorsal  margin  of  basal  ring;  a  transverse  row  of  three  setae 
on  posterior  ring,  lying  ventrad  and  slightly  caudad  of  spiracle  (as  in  E.  spinosa);  tergum  wit* 
basal  ring  unarmed,  posterior  ring  with  a  subterminal  row  of  sharp  black  spines;  two  long 
setae  at  each  end  of  row  and  a  few  small  setae  at  intervals  along  row;  a  solitary  seta  on  basal 
lateral  part  of  posterior  ring;  armature  weaker  on  posterior  segments,  on  segment  7  b 
reduced  to  four  separated  groups  of  setae,  the  outer  groups  wfth  a  single  spine;  sternum  v 
basal  ring  unarmed,  posterior  ring  with  a  subterminal  transverse  row  of  stout  black  spin 
with  two  long  setae  at  each  end  of  row  and  an  isolated  seta  laterad  and  caudad  of  end  of 
row;  at  base  of  ring  on  either  side  a  group  of  two  setae,  the  lateral  one  the  smaller.     Male 
cauda  (Plate  LIII,  260)  with  the  sharp  dorsal  lobes  directed  dorsad,  rather  acute  at  tips,  two 


888  CHARLES  PAUL  ALEXANDER 

setae  on  outer  face  before  tips;  viewed  from  above,  these  setae  seen  to  be  separated  by  a 
deep  U-shaped  notch;  eighth  segment  with  a  dorsal  pentagon  of  five  closely  approximated 
lobes;  just  laterad  of  these  a  group  of  three  setae,  the  posterior  one  long  and  slender,  the 
anterior  one  short  and  stout;  pleural  region  produced  into  a  long  lobe  tipped  with  a  slender 
seta;  on  sternum  two  small  setae  on  either  side.  Female  cauda  (Plate  LIII,  261)  elongate, 
tergal  valves  very  long  and  slender,  two  delicate  setae  on  either  side  before  tip  and  a  stouter 
one  at  tip. 

Nepionotype. —  Ithaca,  New  York,  April  28,  1917. 
Neanotype—  Fall  Creek,  Ithaca,  May  18,  1917 
Paratypes. —  Larvae  and  pupae  with  types. 

Eriocera  longicornis  (Walk.) 

1848    Anisomera  longicornis  Walk.     List  Dipt.  Brit.  Mus.,  vol.  1,  p.  82. 
1869     Eriocera  longicornis  O.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  253-254. 

Eriocera  longicornis  is  probably  the  commonest  species  of  the  genus 
in  the  eastern  United  States.  The  adult  flies  are  sometimes  very  abundant, 
occurring  in  swarms  in  late  afternoon  and  early  evening  in  May,  some 
of  the  swarms  numbering  thousands  of  individuals.  At  other  times  of 
the  day,  the  flies  may  be  found  resting  quietly  on  bushes.  The  larvae 
live  in  the  sand  near  the  water's  edge.  The  pupal  duration  is  seven  days. 
The  detailed  life  history  of  this  species  is  given  on  pages  704  to  708. 

Larva. —  Length,  17-19  mm. 
Diameter,  2-2.3  mm. 

Color,  greenish  brown. 

Body  covered  with  a  long,  appressed,  dark  pubescence.  Penultimate  segment  of  abdomen 
capable  of  great  distention  and  destitute  of  pubescence;  last  segment  of  body  conspicuously 
narrowed.  Spiracular  disk  (Plate  LIV,  265)  surrounded  by  four  slender  lobes,  the  ventral 
pair  the  longer,  bearing  at  tip  elongate  hairs,  some  of  which  exceed  the  lobes  in  length;  on 
lateral  face  at  about  midlength  a  small  pencil  of  hairs;  on  ventral  face  one  or  two  long  setae; 
inner  face  of  ventral  lobes  lined  with  pale  brown,  beginning  as  a  narrow  brown  mark  at  tip, 
at  about  midlength  gradually  expanded  into  an  elongate  triangular  mark,  the  two  lines 
inclosing  between  their  inner  ends  a  pale  linear  mark;  lateral  lobes  similar  to  ventral  lobes, 
fringed  with  long  yellow  hairs  which  are  longer  than  the  lobes;  inner  face  of  lateral  lobes  lined 
with  pale  brown.  Spiracles  rather  large,  separated  by  a  distance  equal  to  about  one  and  one- 
half  diameter  of  one.  Anal  gills  pale.  A  few  setae  in  transverse  alinement  on  last  segment 
behind  lateral  lobes.  Two  pairs  of  short  black  setae  behind  gills. 

Head  capsule  and  mouth  parts  very  similar  to  those  of  Hexatoma,  dorsal  plates  of  capsule 
separate  from  each  other,  not  fused  as  in  the  Ulomorphae;  inner  margins  of  dorsal  plates 
straight  and  parallel.  Labral  sclerite  broadly  transverse,  narrowed  at  ends,  lateral  margins 
produced  into  long  lobes  which  are  densely  tufted  with  short,  golden -yellow  hairs.  Median 
lobe  of  epipharyngeal  region  projecting,  provided  with  two  large  sensory  papillae,  one  on 
either  side,  and  a  few  other  setiferous  papillae  near  tip,  surrounded  by  numerous  hairs. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  889 

Antenna  elongate,  a  little  narrower  at  base,  at  tip  with  three  or  four  hyaline,  seta-like  papillae 
which  are  of  various  diameters  and  shorter  than  the  segment,  the  largest  of  these  papillae 
delicately  sculptured  with  transverse  lines.  Mandible  acute,  curved,  at  about  midlength 
with  a  blunt,  flattened  tooth,  this  with  a  smaller  similar  tooth  in  its  axil.  Maxillary  blade 
very  long  and  slender,  about  half  length  of  capsule. 

Pupa.—  Length:  male,  13.2-15.2  mm.;  female,  14-15.4  mm. 
Width,  d.-s.:  male,  2.1-2.2  mm.;  female,  1.8-1.9  mm. 
Depth,  d.-v.:  male,  2.1-2.3  mm.;  female,  2.2  mm. 

Young  pupae  very  pale;  soft  abdomen  almost  white;  chitinized  anterior  part  of  body  very 
pale  brown.  Older  pupae  much  darker,  the  chitinized  part  becoming  black  with  a  bronzy 
reflection;  abdomen  very  dark  brownish  gray;  breathing  horns  dark  brown  on  apical  half. 

Cephalic  crest  (Plate  LV,  275)  very  prominent,  elongate,  tapering  to  the  subacute  tips; 
lobes  with  blunt  tubercles  behind,  as  well  as  four  long  setae  on  each  lobe,  three  on  dorsal 
margin  and  a  longer  one  on  lateral  face  at  about  midlength;  viewed  from  in  front,  lobes 
separated  by  a  broad,  square  or  U-shaped  notch;  ventral  part  of  crest  produced  forward 
between  antennal  bases  as  a  depressed  lobe  bearing  a  stout  seta  on  either  side.  Tubercle 
on  antennal  scape  very  prominent.  A  slightly  smaller  tubercle  on  either  side  of  clypeus, 
with  a  small  rounded  knob  cephalad  of  each.  Labrum  truncated.  Labial  lobes  roughly 
diamond-shaped.  Maxillary  palpi  very  broad,  rectangular,  tips  truncated.  Antennal 
sheaths  of  male  greatly  elongated,  enlarged  at  base;  viewed  from  beneath,  the  swollen  bases 
nearly  contiguous  on  median  line,  just  above  and  proximad  of  inner  margin  of  eye,  with 
scapal  tubercle  described  above.  Antenna  of  male  exceeding  wing  pads,  those  of  female 
ending  just  beyond  wing  base.  Pronotal  breathing  horns  short  and  stout,  straight,  trans- 
versely wrinkled,  directed  cephalad,  dorsad,  and  laterad;  when  viewed  from  beneath,  com- 
pletely concealed  by  large  cephalic  crest.  Thoracic  notum  convex;  mesonotum  transversely 
wrinkled  (Plate  LV,  277);  median  lobe  of  mesonotal  scutellum  projecting  dorsad  and  cau- 
dad  as  a  blunt  point  (Plate  LV,  272).  Two  or  three  setae  above  wing  axil.  Lateral  angles 
of  thorax  subacute,  with  a  weak  seta.  Wing  sheaths  attaining  end  of  second  abdominal 
segment.  Leg  sheaths  ending  before  caudal  margin  of  third  abdominal  segment;  tarsi  of 
hind  legs  the  longest,  the  two  inner  pairs  ending  about  on  a  level  (Plate  LV,  273). 

Abdominal  segments  (Plate  LVI,  285)  divided  into  a  basal  and  a  posterior  ring;  tergites 
on  posterior  ring  with  a  subterminal  transverse  row  of  spines,  these  varying  from  about  thirty- 
two  on  segment  3  to  about  fourteen  on  segment  7;  these  rows  of  spines  interrupted  on  dorso- 
median  line;  at  each  end  of  row,  three  long  setae,  and  two  additional  groups  of  setae  inter- 
spersed along  row;  two  setae  on  either  side  at  anterior-lateral  angle  of  ring;  tergites  on  basal 
ring  unarmed;  sternites  on  posterior  ring  with  a  subterminal  transverse  row  of  from  twenty- 
four  to  thirty- two  spines,  with  two  setae  at  each  end  of  row;  an  isolated  seta  on  caudo- 
lateral  margin,  close  to  pleura;  a  group  of  two  approximated  setae  near  base  of  posterior 
ring,  on  either  side,  about  at  level  of  spiracles.  Sternites  on  basal  ring  unarmed;  pleuritos 
on  basal  ring  with  a  solitary  seta  at  about  midlength,  but  slightly  nearer  dorsal  margin; 
posterior  ring  with  two  setae  dorso-caudad  of  spiracle,  and  a  third  seta  ventrad  of  it.  Mah 
cauda  (Plate  LVI,  286)  very  blunt,  much  narrower  than  remainder  of  abdomen;  ventral 
lobes  very  blunt;  dorsal  lobes  short,  stout,  ending  in  sharp  points  directed  dorsad,  on  out'*' 
face  a  long  and  a  short  seta;  segment  8  with  a  dorsal  trapezoid  of  four  lobes,  the  posterior 


890  CHARLES  PAUL  ALEXANDER 

pair  the  longer,  each  with  two  setae;  anterior  pair  of  lobes  a  little  more  widely  separated; 
laterad  of  latter  pair  of  lobes,  a  tubercle  bearing  three  setae;  a  long,  powerful  seta  on  pleura; 
two  setae  on  either  side  of  median  line  of  sternum.  Female  cauda  (Plate  LV,  274)  similar, 
but  ventral  lobes  more  pointed,  slightly  exceeding  level  of  dorsal  lobes. 

Nepionotype  —  Ithaca,  New  York,  April  18,  1917. 
Neanotype.—  Fall  Creek,  Ithaca,  May  2,  1913. 
Paratypes. —  Several  hundred  larvae  and  pupae  with  types. 

Eriocera  fultonensis  Alex. 

1912    Eriocera  fultonensis  Alex.    Psyche,  vol.  19,  p.  168-169,  pi.  13,  fig.  7. 

Eriocera  fultonensis  is  a  rather  common  but  usually  local  species  thruout 
the  northeastern  United  States.  The  larvae  are  found  in  the  same  situa- 
tions as  are  described  for  the  other  species  of  the  genus,  in  sand  or  gravel 
near  the  margins  of  usually  large  streams.  On  May  30,  1913,  larvae  were 
found  in  considerable  numbers  along  the  banks  of  Fall  Creek,  Ithaca, 
New  York,  where  they  occurred  in  company  with  numerous  larvae  and 
pupae  of  E.  spinosa,  a  few  large  tabanid  larvae,  a  small  tabanid  pupa, 
and  the  following  beetle  associates:  Bembidion,  Schizogenius,  Tachys, 
Gastrolobium,  and  a  few  others.  The  pupal  duration  is  seven  days  (from 
May  31  to  June  6,  1913). 

Larva.—  Length,  18-26  mm. 
Diameter,  2-2.3  mm. 

Color,  pale  flesh  yellow;  anterior  segments  of  body  a  little  darker. 

Body  long  and  slender.  Spiracular  disk  (Plate  LI  V,  266)  with  ventral  lobes  long  and  slender, 
lateral  lobes  shorter;  ventral  lobes  at  their  tips  with  one  or  two  very  elongate  blackish  hairs 
which  are  from  two  to  three  times  length  of  lobes;  in  addition  to  these  the  usual  apical  fringe 
of  yellowish  hairs  not  exceeding  lobes;  near  base  on  outer  side  a  small  pencil  of  hairs;  each 
ventral  lobe  with  a  very  delicate  capillary  black  line  which  expands  abruptly  at  its  inner 
end  into  a  brown  area,  these  two  areas  inclosing  between  their  proximal  ends  a  more  or  less 
oval  pale  area  (in  some  specimens  the  inner  ends  completely  encircling  this  pale  area,  while 
in  others  the  brown  lines  are  not  continuous  over  the  disk);  Iat3ral  lobes  shorter,  similarly 
fringed  with  yellow  hairs  which  are  longer  than  lobes;  inner  face  of  lobes  with  a  capillary 
black  line.  Spiracles  rounded  oval,  widely  separated. 

Head  capsule  and  mouth  parts  almost  as  in  Hexatoma  megacera  and  Eriocera  longitorms, 
as  already  described;  epipharyngeal  region  (Plate  LIV,  268)  produced  into  a  hemispherical 
rounded  lobe  which  is  densely  covered  with  fine  hairs;  two  large  papillae  on  either  side  near 
tip,  between  them  a  terminal  tuft  of  long  yellow  hairs  surrounding  two  long,  slender,  setif- 
erous  papillae.  Antenna  with  sensory  papillae  at  tip  short,  about  one-third  length  ci'  segment. 

Pupa. —  Length,  14-16  mm. 

Width,  d.-s.,  1.8-2  mm. 
Depth,  d.-v.,  2.5-2.7  mm. 


THE  CKANE-FLIES  OF  NEW  YORK  —  PART  II  891 

Fully  colored  pupae  dark  brown;  cephalic  crest  paler;  pronotal  breathing  horns  pale  yellow, 
darkening  into  brown  at  tips;  wing  pads  light  yellow,  with  dark  venation  showing  clearly; 
pleurites  of  abdomen  of  a  darker  brown  than  sternites  or  tergites. 

Body  somewhat  similar  to  that  of  E.  longicornis,  but  general  form  much  stouter.  Cephalic 
crest  (Plate  LVI,  287)  prominent,  tuberculate,  consisting  of  four  lobes,  the  posterior  lobes 
somewhat  the  larger,  on  posterior  and  lateral  faces  with  two  stout  setae;  a  stout  seta  on 
ventral  face  of  anterior  lobes.  Scapal  spine  lacking.  Clypeal  tubercles  large,  blunt,  with 
a  small  setiferous  tubercle  above  each.  Labrum  bluntly  rounded  at  apex.  Labial  lobes 
elongate,  diamond-shaped,  tips  rather  acute.  Pronotal  breathing  horns  rather  long  and 
slender,  transversely  wrinkled,  longer  than  cephalic  crest.  Mesonotum  (Plate  LV,  279) 
more  convex  than  in  E.  longicornis.  Wing  pads  usually  showing  venation  clearly  on  pale 
background;  vein  r  connecting  Ri  with  Rz+s  distinctive  of  this  species,  lack  of  cell  Mi 
separating  this  pupa  from  that  of  E.  spinosa  and  E.  brachycera.  Legs  sheaths  ending  about 
on  a  level,  the  hind  tarsi  a  little  longer  than  the  two  inner  pairs. 

Arrangement  of  setae  on  abdomen  (Plate  LVI,  288)  about  as  in  E.  longicornis.  Pleura 
with  two  stout  setae  dorsad  and  caudad  of  each  spiracle,  and  a  weak  seta  ventrad  of  spiracle 
and  close  to  it;  basal  ring  with  a  single  pleural  seta;  spicules  on  caudal  margin  of  posterior 
ring  small  and  numerous,  on  intermediate  segments  about  forty  in  number;  seventh  sternite 
with  about  four  to  six  spines  at  each  end  of  row,  the  broad  median  area  devoid  of  spines. 
Female  cauda  (Plate  LV,  276)  with  tergal  valves  exceeding  the  long  sternal  valves, 
scarcely  directed  dorsad  (this  condition  may  be  compared  with  that  in  E.  longicornis).  Male 
caada  with  abdomen  bluntly  rounded  at  tip. 

Nspiinotype.—  Ithaca,  New  York,  May  30,  1913. 
Ne  .mt>/pe  —  With  type  larva,  June  6,  1913. 
Paratypes. —  Larvae  and  pupae  with  types. 

Genus  Penthoptera  Schiner  (Gr.  sorrow  +  wing) 

1.X3     Penthoptera  Schin.     Wien.  Ent.  Monatschr.,  vol.  7,  p.  220. 

Larva.—  Spiracular  disk  surrounded  by  four  blunt  lobes,  the  ventral  pair  a  little  the  longer, 
inner  fac3  not  marked  with  darker,  at  tip  with  one  or  more  long  setae.  Head  capsule  about 
as  in  Eriocera.  Coloration  a  deep  saturated  yellow. 

Pupa.- —  Cephalic  crest  with  lobes  rounded,  setiferous.  No  distinct  tubercles  or  spines 
on  head  or  thorax.  Pronotal  breathing  horns  short,  stout,  cylindrical,  apex  expanded  into 
a  flattened  head,  stem  caarsely  wrinkled,  base  enlarged.  Abdominal  armature  weak. 
Spiracles  not  well  developed. 

.; 

Penthoptera  is  a  small  genus  which  includes  four  European  and  three 
American  species,  two  of  the  latter  occurring  in  tropical  America.  The 
eastern  North  American  Penthoptera  albitarsis,  discussed  below,  has  been 
considered  in  some  detail  by  the  author  in  another  paper  (Alexander, 
1915c: 152-157). 


892  CHARLES  PAUL  ALEXANDER 

Penthoptera  albitarsis  O.  S. 

1869     Penthoptera  albitarsis  0.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  257-258. 

The  larvae  of  Penthoptera  albitarsis  are  usually  not  uncommon  in  rich 
organic  mud  in  shaded  places  thruout  the  range  of  the  species.  Larvae 
of  many  sizes,  some  very  small,  others  apparently  almost  fully  grown, 
may  be  found  at  a  single  time.  This  would  probably  indicate  that  the 
species  emerges  at  intervals  thruout  the  summer,  rather  than  that  it  is 
double-brooded. 

Larva. —  Length,  10-12  mm. 
Diameter,  1-1.2  mm. 

Color  bright  chestnut-yellow,  anterior  half  of  body  richer-  and  deeper-colored;  thoracic 
sogments  suffused  with  brown;  skin  with  a  silky,  iridescent  reflection. 

Body  provided  with  numerous  long,  appressed  hairs.  A  few  setae  on  body,  the  following 
being  the  most  conspicuous:  one  on  lateral  do:  sal  margin  of  last  segment,  near  base  of 
lateral  lobes;  a  series  of  four  groups  of  one  or  two  in  each  row  across  dorsal  surface  of  the 
three  thoracic  segments  at  about  midlength;  a  group  of  two  or  three  long  setae  on  sides  near 
caudal  margin  of  segments.  Subterminal  enlargement  of  abdomen  with  about  twenty-five 
transverse  rows  of  fine  points. 

Spiracular  disk  (Plate  LVII,  292  and  293)  with  four  blunt  lobes;  ventral  lobes  densely 
fringed  with  long,  pale  hairs,  those  toward  ends  of  lobes  longer;  one  or  more  elongate  setae 
near  tip  of  each  ventral  lobe,  these  being  longer  than  lobes  themselves;  lateral  lobes  with 
a  similar  fringe  of  rather  short,  yellow  hairs;  spiracular  disk  almost  free  from  dark  mark- 
ings, a  pale  brown  line  extending  dorsad  from  each  spiracle  and  an  indistinct  brownish  line 
along  ventral  margin  of  lateral  lobes.  Spiracles  circular.  Anal  gills  four,  pale.  Head 
capsule  rather  broad,  dorsal  plate  with  inner  anterior  angles  rounded.  Labrum  (Plate 
LVII,  289)  almost  as  in  Eriocera  spinosa,  the  extreme  cephalic  epipharyngeal  parts  with  the 
usual  papillae  and  seti'erous  tubercles;  lateral  papillae  bearing  at  their  tips  three  or  four 
slender  pegs;  between  these  papillae  two  pairs  of  setiferous  tubercles,  a  basal  larger  pair 
and  a  more  apical  smaller  pair.  Mandible  (Plate  LVII,  291)  a  little  more  curved  than  is 
usual  in  this  subtribe,  inner  margin  with  a  double  tooth  at  about  midlength. 

Pupa.—  Length,  10-10.5  mm. 

Width,  d.-s.,  1.4-1.5  mm. 
Depth,  d.-v.,  1.5-1.6  mm. 

Thorax  dark  brown,  wing  and  leg  sheaths  paler;  pronotal  breathing  horns  dark  brown, 
swollen  bases  and  tips  much  paler,  light  orange;  abdomen  brownish  yellow. 

Cephalic  crest  consisting  of  two  widely  separated  rounded  lobes  behind,  each  bearing  two 
setae;  anteriorly  the  crest  appearing  as  a  large  depressed  lobe  between  antennal  bases,  with 
a  very  large,  stiff  seta  on  either  side.  A  powerful  seta  on  each  side  of  region  of  clypeus.  A 
seta  on  genal  region  between  eye  and  sheath  of  maxillary  palpus.  Labrum  evenly  rounded 
or  a  little  truncated  at  apex.  Labial  lobes  widely  separated,  roughly  rounded  or  indistinctly 
pentagonal  in  outline.  Maxillary  palpi  broad,  ending  bluntly  beneath  or  just  before  antennal 
sheaths.  Antennae  ending  just  beyond  wing  root  in  female,  considerably  longer  in  male. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  893 

Pronotal  breathing  horns  (Plate  LVII,  294)  short,  stout,  cylindrical,  apex  expanded  into  a 
flattened  circular  head,  stem  coarsely  and  transversely  wrinkled,  base  enlarged;  breathing 
horns  widely  separated  at  their  bases,  but  bent  proximad  so  as  to  be  almost  contiguous  at  their 
tips;  two  strong  setae  laterad  of  base  of  breathing  horn  and  an  additional  one  in  front  of  it. 
Mesonotum  transversely  wrinkled,  with  a  distinct  carina  anteriorly  (Plate  LVIII,296).  Two 
groups  of  two  setae  on  either  side  of  median  line,  with  an  additional  solitary  seta;  two  longer 
setae  above  wing  axil.  Wing  sheaths  ending  before  tip  of  second  abdominal  segment.  Leg 
sheat'is  ending  at  from  two-thirds  length  to  opposite  end  of  third  abdominal  segment;  tarsal 
sheaths  ending  about  on  a  level,  or,  in  some  specimens,  the  hind  tarsi  a  little  longer  than  the 
others  (Plate  LVIII,  297). 

Abdomen  indistinctly  divided  into  a  narrow  basal  ring  and  a  broader  posterior  ring;  basal 
ring  further  very  indistinctly  subdivided  into  two  subequal  annuli.  Abdominal  armature 
weak;  on  stern ites  a  subterminal  transverse  row  of  delicate  spines  with  two  setiferous  tubercles 
at  each  end  of  row;  on  posterior  ring  two  setae  on  either  side  at  about  midlength;  tergites 
with  four  groups  of  two  or  three  setae  near  posterior  margin  and  an  additional  group  of  two 
setae  on  lateral  margin  of  posterior  ring  near  base;  pleural  region  with  a  stiff  seta  on  extreme 
anterior  part  of  basal  ring.  Spiracles  distinct;  a  group  of  two  setae  caudad  and  slightly 
ventrad  of  each  spiracle,  with  an  additional  solitary  seta  caudo-ventrad  of  these.  Female 
cauda  (Plate  LVIII,  298)  with  tergal  valves  of  ovipositor  only  a  little  longer  than  sternal 
valves,  at  tip  ending  in  a  short,  rather  blunt  point  directed  dorsad;  on  outer  face  before  tip 
a  short,  stiff  seta;  segment  8  on  dorsum  with  a  close  trapezoid  of  four  irregular  lobes;  two 
setiferous  tubercles  on  dorsal  and  lateral  part  of  eighth  segment,  the  more  dorsal  of  these 
with  two  setae,  the  lateral  one  with  a  single  seta;  sternum  with  four  stout  setae,  of  which  two 
are  lateral  and  two  are  median  in  position.  Male  cauda  (Plate  LVII,  295.  and  Plate 
LVIII,  299)  with  sternal  valves  short  and  blunt;  tergal  valves  slender,  ending  in  an  acute 
point  directed  dorsad;  a  few  short  setae  on  outer  face  before  tip. 

Nepionotype. —  Ithaca,  Now  York,  May  25,  1917. 
Neanotype. —  Bool's  hillside,  Ithaci,  June  5,  1917. 
Paratypes. —  Abundant  larvae  and  a  few  pupae  with  types. 

Subtribe  Polymeraria 

Genus  Polymera  Wiecbmann  (Gr.  many  +  part) 
1821     Polymera  Wied.     Dipt.  Exot.,  vol.  1,  p.  40. 

Polymera  is  a  tropical  American  genus  including  fifteen  described 
species,  one  of  which,  Polymera  georgiae  Alex.,  occurs  in  the  southeastern 
United  States.  A  single  additional  species,  P.  magnified  Meunier  (1906: 
385),  has  been  described  from  the  Baltic  amber  (Lower  Oligocene).  The 
only  species  concerning  the  ecology  of  which  we  have  any  record  is  P. 
geniculata  Alex.,  which  has  been  found  living  in  crabholes  beneath  rocks 
in  Porto  Rico.  In  this  connection  the  long-horned  deinoceritine  mosquitoes 


894  CHARLES  PAUL  ALEXANDER 

which  live  in  similar  habitats  should  be  considered.     Howard,  Dyar,  and 
Knab  (1915:213)  say,  in  describing  these  mosquitoes: 

These  crab-hole  inhabiting  species  possess  peculiarly  developed  antennae  in  order,  as  we 
suppose,  to  enable  them  to  detect  the  approach  of  their  crustacean  host  and  fly  out  of  the 
hoi ?s  before  being  overwhelmed  in  the  water  in  the  bottom  by  the  incursion  of  the  crab, 
whose  body  must  completely  fill  the  entrance  to  the  hole. 

It  is  curious  and  suggestive  that  the  males  of  Polymera  should  likewise 
possess  elongated  and  very  complicated  antennae. 

Tribe  Pediciini 

The  Pediciini  constitutes  a  small  tribe  which  seems  to  be  divisible  into 
two  well-marked  subtribes,  the  more  generalized  Adelphomyaria  indicating 
a  relationship  with  the  Hexatomini. 

The  larvae  of  the  Pediciini  have  the  labrum  broad  and  the  epipharynx 
usually  feebly  armed.  The  mentum  is  completely  divided  into  two  parts, 
each  half  with  not  more  than  four,  usually  three,  teeth.  The  hypopharynx 
is  labriform.  The  maxilla  consists  of  two  lobes,  distinct  and  separate 
in  the  Adelphomyaria,  more  or  less  approximated  or  fused  in  the  Pedi- 
caria.  The  mandible  is  powerful,  ending  in  a  strong  apical  point;  the 
cutting  edge  has  about  four  teeth;  there  is  a  simple  tuft  of  setae  on  the 
prosthecal  region  in  the  Dicranotae  and  in  Pedicia.  The  head  capsule 
is  very  elongate,  massive,  and  compact,  with  the  posterior  incisions  very 
shallow.  In  the  Adelphomyaria  the  cauda  is  surrounded  by  four  lobes 
which  are  fringed  with  exceedingly  elongate  hairs;  in  the  Pedicaria  there 
are  two  ventral  caudal  lobes,  each  tipped  with  a  very  few  setae.  The 
anal  gills  are  four  in  number  and  are  segmented,  the  terminal  segment 
being  more  or  less  retractile.  In  the  Pedicaria  prolegs  are  developed 
on  the  abdominal  segments  of  some  of  the  genera. 

All  of  the  species  of  the  tribe,  so  far  as  is  known  to  the  writer,  are 
carnivorous  in  their  larval  state,  which  is  spent  in  mud  or  earth  close  to 
water. 

The  two  subtribes  of  the  Pediciini  may  be  separated  as  follows: 

Larvae 

Spiracular  lobes  four  in  number,  fringed  with  very  long,  delicate  hairs. 

Adelphomyaria  (p.  895) 

Spiracular  lobes  two  hi  number,  ventral  in  position,  each  tipped  with  from  six  to  eight  setae. 

Pedicaria  (p.  899) 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  895 

The  most  important  literature  on  the  tribe  Pediciini  is  as  follows: 

Pedicia  rivosa General Scheffer,  in  Rossi,  1848:9. 

Pedicia  rivosa Larva,  pupa,  general .  .  Beling,  1879:45-46. 

Pedicia  rivosa General Reuter,  1893. 

Pedicia  rivosa General Wesenberg-Lund,  1915: 335. 

Pedicii  albivitta Larva NeecT.iam,  1903: 285-286;  1905: 8. 

Tricyphoqi  immaculata Larva,  general Beling,  1879:47. 

Tricyphona  immaculata Larva,  pupa,  general.  .  De  Meijere,  1916: 195-196. 

Tricyphona  schineri Larva,  pupa,  general.  .  Beling,  1879:47. 

Dicranota  bimaculata Larva,  pupa,  general.  .  Miall,  1893. 

Dicranota  bimaculata Larva,  pupa Griinberg,  1910:66-67.    (Copy.) 

Dicranota  bimaculata General Wesenberg-Lund,  1915: 342-343. 

Dicranota  bimaculata Larva .- Malloch,  1915-17  b:  219-220. 

(Copy.) 

Rhaphidolabis  tenuipes Larva,  general Needham,  1908 a: 212-214. 

Subtribe  Adelphomyaria 

The  division  Adelphomyaria,  as  known,  includes  but  the  single  genus 
Adelphomyia  Bergroth,  a  curious  genus  of  small  crane-flies  which,  in  the 
general  appearance  of  the  adults,  strongly  suggest  the  hexatomine  sub- 
tribe  Limnophilaria.  The  immature  stages  have  not  been  associated  with 
the  adult  flies  by  rearing,  and  there  is,  of  course,  the  possibility  of  a 
mistaken  reference.  The  immature  stages  of  the  insect  herein  described 
are  easily  recognized,  however,  and,  no  matter  to  what  group  it  belongs, 
it  deserves  subtribal  rank  under  the  Pediciini. 

Genus  Adelphomyia  Bergroth  (Gr.  brother  +  fly) 

1891     Adelphomyia  Bergr.    Mittheil.  Naturf.  Ges.  Bern,  1890,  p.  134. 

Larva  (supposition).- — Body  with  pencils  of  stiff  setae,  producing  a  spiny  appearance 
Spiracular  disk  surrounded  by  four  short  lobes  which  are  fringed  with  exceedingly  elongate 
hairs.  Spiracles  large,  separated  by  a  distance  less  than  the  diameter  of  one.  Head  cap- 
sule long  and  massive,  all  the  plates  firmly  united  except  behind.  Mandible  acutely  pointed. 
Maxilla  of  two  elongate  separated  lobes.  Antenna  two-segmented,  the  terminal  segment 
with  three  small  papillae.  Hypopharynx  labriform.  Mentum  completely  divided,  each 
half  with  four  teeth,  the  middle  pair  on  each  side  the  largest  (minuta,  supposition)  or  the 
second  from  the  inside  the  largest  (americana,  supposition). 

Pupa  (supposition^ .—  Cephalic  crest  small,  each  lobe  with  three  setiferous  tubercles. 
Pronotal  breathing  horns  moderate  in  length,  broadly  tipped  with  light  yellow.  Mesonotum 
unarmed.  Wing  sheaths  extending  beyond  base  of  third  abdominal  segment.  Leg  sheaths 
extending  to  beyond  base  of  fifth  abdominal  segment.  Abdominal  armature  weak,  especially 
on  posterior  segments. 

Adelphomyia  is  a  small  genus  of  crane-flies,  including  but  four  European 
and  three  North  American  species,  and  a  doubtful  species  from  Africa.  The 


896  CHARLES  PAUL  ALEXANDER 

insects  resemble  tiny  species  of  the  genus  Limnophila.  The  adult  flies 
are  not  uncommon  on  rank  herbage,  especially  ferns,  in  woods  and  usually 
near  running  water.  As  already  stated,  the  immature  stages  have  not 
been  reared,  but  larvae  found  by  the  writer  in  Maine  are  referred  with 
considerable  confidence  to  Adelphomyia  americana  and  A.  cayuga,  while 
larvae  and  pupae  of  another  species  taken  at  Ithaca,  New  York,  seem  to 
be  those  of  A.  minuta.  The  larvae  show  a  curious  combination  of  tribal 
characters.  The  general  appearance  and  the  structure  of  the  spiracular 
disk  are  altogether  those  of  one  of  the  Hexatomini;  but  the  head  capsule 
and  the  details  of  the  mouth  parts  indicate  a  relationship  with  the  Pedi- 
ciini  that  cannot  be  denied. 

Adelphomyia  minuta  Alex,  (supposition) 

1911     Adelphomyia  minuta  Alex.     Can.  Ent.,  vol.  43,  p.  287-288. 

Adelphomyia  minuta  is  a  characteristic  late  spring  species,  common  in 
boggy  woods  and  on  vegetation  along  rapid  streams.  Larvae  and  pupae 
which  are  referred  to  this  species  were  sifted  from  organic  mud  taken  on 
Bool's  hillside,  Ithaca,  New  York,  thruout  May  and  early  June,  1917. 
The  pupae  referred  to  this  species  strongly  resemble  those  of  Dicrano- 
phragma  but  in  reality  are  very  different.  The  species  is  discussed 
herewith  in  the  hope  that  it  may  be  definitely  recognized  in  the  future. 

Larva. —  Length,  4.5-5  mm. ;  caudal  fringe,  2  mm.  additional. 
Diameter,  0.4  mm. 

Coloration,  saturated  yellow  with  a  faint  orange-brown  tinge. 

Form  narrow,  body  tapering  gradually  to  both  ends,  spiracular  disk  narrowed.  Body 
clothed  with  a  delicate  appressed  pubescence  and  numerous  tufts  of  conspicuous  stiff  hairs 
which  produce  a  spiny  or  bristly  appearance;  the  more  conspicuous  of  these  tufts  located  on 
pleural  region,  there  being  three  such  rows  on  abdominal  segments  • —  one  on  basal  ring,  the 
second  and  largest  at  base  of  posterior  ring,  and  the  third  just  before  posterior  margin  of 
segment  and  more  ventral  in  position;  only  the  large  intermediate  tuft  present  on  thoracic 
segments,  the  small  brush  on  anterior  annulus  of  abdominal  segments  lacking.  Spiracular 
disk  (Plate LIX,  305)  with  four  lobes;  ventral  pair  not  more  than  three  times  length  of  lateral 
pair;  inner  face  of  lobes  margined  with  brown;  at  tip  of  ventral  lobes  a  fringe  of  exceedingly 
elongate  hairs,  which  are  from  one-third  to  nearly  one-half  length  of  entire  body  and  about 
fifteen  times  length  of  lobes  bearing  them;  a  stiff  sensory  bristle  located  in  black  margin 
at  tip  of  lobe;  lateral  lobes  with  fringe  of  hairs  relatively  much  shorter,  tho  still  long. 
Spiracles  large,  close  together,  separated  by  a  distance  less  than  diameter  of  one,  the  middle 
piece  black,  the  ring  pale  yellow.  Anal  gills  four,  slender,  hyaline,  each  subdivided  by  con- 
strictions into  four  lobes  which  are  gradually  narrowed  from  the  base  outward,  the  last  being 
cylindrical. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  897 

Head  capsule  very  compact,  lateral  plates  united  with  broad  dorsal  plate  except  for  a  short 
distance  behind.  Labrum  (Plate  LIX,  300)  broadly  transverse;  cephalic  margin  truncate  or 
very  slightly  concave;  epipharyngeal  region  with  about  five  transverse  interrupted  rows  of 
setae.  Mentum  (Plate  LIX,  302)  of  two  entirely  separated  plates,  each  half  with  four  teeth, 
those  at  the  ends  the  smallest,  the  outermost  tooth  tending  to  be  reduced,  the  two  middle 
teeth  of  each  side  notably  larger  than  the  others;  outside  of  mentum  a  thin  plate,  its  inner 
proximal  margins  nearly  contiguous  at  median  line  of  body.  Hypopharynx  labriform,  pro- 
jecting beyond  level  of  both  labrum  and  mentum;  outer  lateral  angles  densely  hairy, 
median  posterior  area  with  about  eight  small,  hyaline  spines.  Antenna  (Plate  LIX,  301),  in 
caustic-potash  mounts,  hyaline;  basal  segment  elongate-cylindrical,  bearing  at  its  tip  about 
two  papillae  as  follows:  a  long,  slightly  curved,  cylindrical  papilla  which  is  finely  sculptured, 
and  immediately  proximad  of  this  a  slightly  smaller  second  segment  of  the  antenna,  bearing 
near  its  tip  three  tiny  papillae,  an  inner  flattened  subspatulate  blade,  and  two  longer  cylin- 
drical papillae;  near  base  of  this  second antennal  segment  a  long  seta.  Mandible  (Plate  LIX, 
303)  rather  long  and  slender,  apical  point  narrow;  about  four  small  lateral  teeth  near  ventral 
cutting  edge,  the  most  basad  of  these  acute;  a  large  dorsal  tooth  on  cutting  edge  and  two 
very  small  acute  teeth  near  base;  on  dorsal  face  of  mandible  two  long  setae,  and  two 
somewhat  shorter  setae  near  heel  of  mandible;  about  five  or  six  stout  setae  at  prosthecal 
region.  Maxilla  (Plate  LIX,  304)  consisting  of  two  elongated  lobes  which  are  separate  from 
each  other,  the  outermost  the  longer,  the  pale  rounded  palpus  terminal  in  position;  inner 
lobe  shorter,  with  three  elongate  setae,  of  which  one  is  apical  and  the  longest  is  sub-basal  in 
position. 

Pupa. —  Length,  4.5-4.7  mm. 
Depth,  d.-v.,  0.8  mm. 

Color  light  yellowish  brown,  the  thoracic  dorsum  paler;  pronotal  breathing  horns  dark 
brown,  apical  quarter  abruptly  light  yellow;  abdomen  brown,  lateral  and  posterior  parts 
of  each  segment  darker. 

Cephalic  crest  small,  each  lobe  with  three  small  setiferous  punctures  on  anterior  face; 
ventrad  of  crest  between  antennal  bases  a  prominent  median  lobe.  Labrum  broad;  apex 
truncated,  indistinctly  bifid.  Labial  lobes  large,  with  a  deep  U-shaped  median  notch  behind. 
Maxillary  palpi  rather  short  and  stout,  ending  before  joint  of  fore  legs  (Plate  LX,  307). 
Pronotal  breathing  horns  moderately  elongate,  curved  slightly  laterad,  strongly  divergent, 
cylindrical,  of  uniform  diameter  thruout  their  length.  Mesonotum  not  very  gibbous  (Plate 
LX,  306).  Thorax  with  a  high  anterior  median  carina.  Wing  sheaths  extending  beyond 
base  of  third  abdominal  segment,  the  venation  indistinct.  Leg  sheaths  rather  long,  ending 
at  about  one-third  length  of  fifth  abdominal  segment;  hind  legs  much  longer  than  the 
others,  middle  legs  a  very  little  longer  than  fore  legs. 

Abdominal  segments  indistinctly  subdivided  into  two  rings,  a  narrow  anterior  ring  and  a 
much  broader  posterior  ring;  abdominal  armature  very  weak,  lacking  on  segment  7;  basal 
annulus  on  tergites  and  sternites  with  a  number  of  small  pits  on  sides,  these  sometimes  sparse 
or  lacking;  posterior  ring  with  a  transverse  row  of  numerous  long,  stout  setae  or  delicate 
spines  before  margin.  Spiracles  weak,  at  base  of  posterior  ring.  Male  cauda  (Plate  LX, 
309  and  310)  with  sternal  lobes  elongate,  contiguous  except  at  extreme  tips;  tergal  lobes 
terminating  in  very  slender  and  acute  curved  points,  directed  caudad  and  dorsad;  tergite 


898  CHAKLES  PAUL  ALEXANDER 

8  swollen,  with  four  blunt  lobes,  the  posterior  pair  large,  with  their  posterior  faces  setiferous. 
Female  cauda  (Plate  LX,  308)  with  sternal  valves  much  shorter  than  tergal  valves  and  rather 
blunt  at  tips;  tergal  valves  broad  at  base,  narrowed  at  tips,  which  terminate  in  acute  black 
spines,  situated  on  lateral  margin  before  apex,  the  spines  directed  dorsad,  laterad,  and  caudad. 

The  larvae  were  common  near  Ithaca,  New  York,  from  May  10  to 
June  5,  1917.  The  pupae  are  described  from  one  male  and  two  females 
washed  from  mud  from  Bool's  hillside,  at  Ithaca,  on  June  11,  1917.  A 
little  later  in  June  adult  flies  of  this  species  were  common  at  this  location. 

Adelphomyia  americana  Alex,  (supposition) 

1912    Adelphomyia  americana  Alex.     Pomona  Journ.  Ent.,  vol.  4,  p.  829-831. 

Larvae  that  were  rather  common  in  the  rich  organic  mud  from  the 
Standpipe  Woods,  Orono,  Maine,  from  July  1  to  14,  1913,  are  referred 
with  some  doubt  to  Adelphomyia  americana.  They  are  unquestionably 
congeneric  with  the  species  last  described  (A.  minuta,  supposition)  and 
with  the  form  discussed  in  the  following  pages  as  A.  cayuga.  A  short 
time  after  the  larvae  of  these  three  species  were  obtained,  the  adult  flies 
appeared  in  considerable  numbers  in  the  same  situations  and  there  seems 
to  be  but  little  doubt  as  to  the  reference.  Associated  with  the  larvae 
of  this  species  in  the  organic  mud  were  a  few  larvae  of  Penthoptera, 
Rhaphidolabis,  and  other  species  of  crane-flies. 

The  larva  of  the  present  species  averages  larger  than  that  of  A .  minuta, 
when  fully  grown  measuring  5.5  millimeters  in  length.  In  coloration 
it  is  light  yellow.  The  ventral  lobes  of  the  spiracular  disk  have  the  dark 
markings  on  their  inner  face  much  more  extensive,  the  apical  half  being 
suffused  with  brown.  The  mouth  parts  are  similar  to  those  of  A.  minuta 
as  described,  but  the  outermost  of  the  two  large  teeth  of  each  half  of  the 
mentum  is  reduced  in  size  so  that  only  a  single  tooth  is  of  conspicuous  size. 

(Described  from  larvae  taken  at  Orono,  Maine.     No.  66-1913.) 

Adelphomyia  cayuga  Alex,  (supposition) 

1912    Adelphomyia  cayuga  Alex.     Pomona  Journ.  Ent.,  vol.  4,  p.  831. 

The  supposed  larva  of  Adelphomyia  cayuga  occurred  with  specimens 
of  the  preceding  (A.  americana,  supposition)  at  Orono,  Maine,  from  July 
1  to  13,  1913.  This  is  a  smaller  species  than  the  preceding,  measuring 
but  4  millimeters  in  length,  and  is  much  paler,  the  color  being  almost 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  899 

white.     The  condition  of  the  mental  teeth  is  almost  as  in  A.  americana, 
but  the  outermost  of  the  two  large  intermediate  teeth  is  a  little  larger. 
The  larvae  were  rather  frequent  in  the  mud  beneath  saturated  moss. 
(Described  from  larvae  taken  at  Orono,  Maine.     Nos.  57-  and  67-1913.) 

Subtribe  Pedicaria 

The  Pedicaria  comprise  a  well-defined  division  of  the  tribe  Pediciini, 
including  two  groups  of  genera  —  the  more  generalized  Pediciae,  with 
the  genera  Pedicia,  Tricyphona,  Ornithodes,  and  Rhaphidolabina,  and 
the  specialized  Dicranotae,  with  the  genera  Dicranota,  Rhaphidolabis, 
and  probably  Polyangaeus. 

The  genera  of  the  Pedicaria  may  be  divided  in  the  main  as  follows: 

Larvae 

1.  Abdomen   without  prolegs,  but  with  raised  welts  on  segments  4  to  7,   these  covered 

with  a  microscopic  scurfiness.     (Group  Pediciae) 2 

Abdomen  with  conspicuous  cylindrical  prolegs  on  segments  3  to  7,  these  with  circlets 
of  conspicuous  chitinized  hooklets  around  their  ends Group  Dicranotae  (p.  906) 

2.  Abdominal  segments  4  to  7  with  raised  welts  on  both  dorsal  and  ventr  t.l  surfaces;  sid^s  of 

hypopharynx  not  parallel,  narrowed  toward  base Rhaphidolabina  Alex.  (p.  901) 

Abdominal  segments  4  to  7  with  welts  on  ventral  surface  only;  sides  of  hypopharynx 
subparallel -. 3 

3.  Size  very  large,  when  fully  grown  40  mm.  in  length;  mental  plates  without  a  small  lateral 

tooth Pedicia  Latr.  (p.  899) 

Size  smaller,  when  fully  grown  under  30  mm.  in  length;  mental  plates  with  a  small  lateral 
tooth < Tricyphona  Zett.  (p.  903) 

The  writer  is  unable  to  separate  the  genera  of  the  group  Dicranotae 
with  the  material  available,  and  is  inclined  to  suspect  the  congenerousness 
of  Rhaphidolabis  with  Dicranota.  The  characters  given  by  Malloch 
(1915-17  b:  217)  to  separate  the  two  genera  do  not  hold  at  all  in  a  series. 

It  is  impossible  also  to  key  the  pupae  with  the  present  knowledge  of 
the  group. 

Group  Pediciae 

Genus  Pedicia  Latreille  (Gr.  a  field ) 

1809     Pedicia  Latr.     Hist.  Nat.  Crust,  et  Ins.,  vol.  4,  p.  255. 

1916     Daimiotipula  Matsumura.     Thous.  Ins.  Japan,  add.  2,  p.  463. 

Pedicia  is  a  small  genus  including  but  six  described  species,  one  occurring 
in  Europe,  one  in  Japan,  and  four  in  North  America.  Of  the  last-named, 


900  CHARLES  PAUL  ALEXANDER 

two  are  eastern  and  two  are  western  in  their  distribution.     They  include 
the  largest  and  most  beautiful  species  of  the  Limnobiinae. 

The  European  Pedicia  rivosa  (Linn.)  was  found  by  Beling  (1879:45-46) 
living  in  brooks  and  springs,  or  in  wet  spots  among  saturated  leaves  and 
other  debris,  sometimes  associated  with  the  larvae  of  Tipula  lutescens 
Fabr.  The  pupae  live  in  cylindrical  vertical  burrows,  clothed  in  the 
last  larval  skin,  and  are  able  to  move  up  and  down  in  these  passages 
Pupation  lasts  from  one  to  two  weeks. 

Needham  (1903:285-286,  and  1905:8)  was  the  first  to  describe  and 
figure  the  larva  of  the  commonest  eastern  species,  Pedicia  albivitta  Walk. 

Pedicia  albivitta  Walk. 

1848     Pedicia  albivitta  Walk.     List  Dipt.  Brit.  Mus.,  vol.  1,  p.  37. 

Pedicia  albivitta  is  a  beautiful  fly,  common  and  widely  distributed 
thruout  the  northeastern  United  States  and  Canada.  The  adults  are 
on  the  wing  in  midsummer,  and  a  few  individuals  may  usually  be  found 
in  June.  The  much  rarer  and  more  local  P.  contermina  Walk,  is  a  vernal 
species,  on  the  wing  in  May  and  early  June. 

The  larvae  of  P.  albivitta  live  in  cold  springs  and  beneath  saturated 
moss  at  the  edge  of  streams.  The  writer  has  never  succeeded  in  rearing 
this  species  to  the  adult  condition. 

Larva. —  Length,  40-44  mm. 
Diameter,  5-5.5  mm. 

Color  dark  grayish  brown  above,  paler  at  sutures  and  on  posterior  half  of  body;  paler 
beneath,  more  grayish. 

Body  covered  with  a  short,  appressed,  dusky  pubescence.  Thoracic  segments  with  a 
pencil  of  small  setae  on  pleural  region.  Abdominal  segments  with  a  few  delicate  lateral 
setae  on  posterior  ring,  at  about  midlength  of  segments.  Ventral  creeping-welts  on  abdominal 
segments  4  to  7  completely  divided  on  median  line,  the  welts  covered  with  a  microscopic 
scurfiness.  Spiracles  (Plate  LXI,  311)  circular,  separated  by  a  distance  about  equal  to 
diameter  of  one,  situated  on  a  slightly  protuberant  elevation.  Spiracular  lobes  two,  ventral 
in  position,  short,  slender,  each  with  about  six  setae  *at  tip.  Anal  gills  (Plate  LXI,  315) 
short,  stout  at  base,  before  tip  a  constriction  cutting  off  the  elongate  conical  terminal 
segment,  which  is  partly  telescopic  within  the  next  basal  segment. 

Head  capsule  (Plate  LXI,  312)  massive,  elongate,  as  in  this  division.  Labrum  broadly 
transverse,  lateral  parts  a  little  enlarged  and  projecting  anteriorly  into  blunt  lobes,  with  a 
long  seta  near  inner  margin;  median  region  of  labrum  with  two  widely  separated  setae,  just 
laterad  of  each  of  which  is  a  small  papilla.  Epipharynx  roughened  into  a  narrow  transverse 
band  of  small  spines.  Mentum  completely  divided,  each  half  continuous  with  ventral  plate 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  901 

of  same  side;  anterior  margin  of  each  half  with  three  slender,  flattened  teeth,  the  middle  one 
of  which  is  slightly  the  shortest.  Hypopharynx  conspicuous,  labriform;  anterior  margin  with 
a  deep  notch  to  form  distinct  lobes  at  lateral  angles,  and  with  several  rows  of  small  chitinized 
tubercles.  Antenna  (Plate  LXI,  313)  small;  basal  segment  elongated,  slightly  curved,  a 
circular  auditory  plate  near  base,  at  tip  with  numerous  papillae,  two  of  which  are  very  long, 
nearly  as  long  as  segment  itself;  in  addition  to  these,  three  or  four  tiny  cylindrical  papillae. 
Mandible  powerful;  ventral  cutting  edge  with  a  row  of  about  four  teeth  which  are  succes- 
sively enlarged  from  tip  to  bass;  basal  tooth  very  broad  and  flat,  with  outer  margin  truncate 
or  slightly  concave;  teeth  on  dorsal- cutting  edge  indistinct;  a  pencil  of  moderately  delicate 
setae  on  scrobal  region  of  mandible,  and  another  at  prosthecal  region.  Maxilla  elongate; 
outer  lobe  larger  than  inner  lobe,  chitinized,  apex  with  a  very  flat  circular  palpus  (Plate 
LXI,  314)  which  is  provided  with  a  few  disklike  papillae  around  margin  and  a  few  scattered 
sensory  papillae  over  pale  apex;  inner  lobe  with  a  long,  powerful  seta  on  ventral  face  and 
smaller  setae  near  tip. 

Nepionotype. —  Ithaca,  New  York,  June  1,  1917. 
Paratypes.—  Larvae  from  type  locality. 

Genus  Rhaphidolabina  Alexander  (Gr.  diminutive  of  Rhaphidolabisfy 
1016    friaphiMabina  Alex.     Proc.  Acad.  Kit.  Sci.  Phih.,  p.  540-541. 

Larva.—  Body  covered  with  an  abundant,  appressed  pubescence  and  tufts  of  erect  hairs 
which  are  more  numerous  on  anterior  end  of  body.  Creeping-welts  on  dorsal  and  ventral 
surfaces  of  abdominal  segments  4  to  7.  Spiracular  lobes  two,  moderately  elongated,  each 
with  about  six  hairs  at  tip.  Spiracles  large.  Anal  gills  four,  long  and  diaphanous.  Head 
capsule  long  and  massive.  Mandible  powerful.  Antenna  short,  with  two  elongate  papillae. 
Hypopharynx  labriform,  anterior  margin  concave.  Mentum  completely  divided,  each  half 
with  three  large  teeth  and  a  much  smaller  lateral  tooth. 

Pupa.—  Labrum  truncate.  Pronotal  breathing  horns  short-cylindrical  or  slightly  flattened. 
Abdominal  segments  with  circular  areas  of  spicules  on  pleurites.  Intermediate  tergites  with 
a  broad  transverse  band  of  tiny  spines. 

The  genus  Rhaphidolabina  includes  only  R.  flaveola,  a  curious  pallid 
fly  of  the  northeastern  United  States,  serving  as  a  connecting  link  between 
Tricyphona  on  the  one  hand  and  the  Dicranotae  on  the  other.  The 
adult  flies  are  common  on  rank  vegetation  in  cold  woods.  The  immature 
stages  are  spent  in  rich  organic  earth  in  the  same  situations  as  are  fre- 
quented by  the  adult  flies. 

Rhaphidolabina  flaveola  (O.  S.) 

1869    Rhaphidolabis  flaveola  0.  S.    Mon.  Dipt.  N.  Amer.,  part  4,  p.  288. 
The    writer  has  found  the  larvae  of  Rhaphidolabina  flaveola  in  Maine 
and  in  New  York.     At  Orono,  Maine,  larvae  were  numerous  in  the  rich 


902  CHARLES  PAUL  ALEXANDER 

organic  mud  of  the  Standpipe  Woods.  One  large  larva  was  placed  in 
a  watch  crystal  with  a  fully  grown  larva  of  Adelphomyia  cayuga  (supposi- 
tion). It  at  once  seized  the  latter  in  its  mandibles  at  about  the  third 
abdominal  segment,  and  carried  it  helplessly  all  around  the  dish, 
occasionally  shaking  it,  quite  as  a  terrier  does  a  rat. 

Larva. —  Length,  9.8  mm. 

Diameter,  0.8  mm. 

Color,  brown  to  orange-yellow  on  anterior  segments  of  body,  becoming  darker  on  abdominal 
segments  due  to  the  increase  of  pubescence. 

Body  covered  with  an  abundant,  appressed  pubescence.  Form  terete;  body  moderately 
elongated,  gradually  narrowed  toward  both  ends.  Abdominal  segments  2  to  8  divided 
into  a  narrow  basal  ring  and  a  much  broader  posterior  ring;  segments  4  to  7  with  conspicuous 
dorsal  and  ventral  welts,  which  are  larger  and  more  conspicuous  on  posterior  segments; 
these  welts  occupying  basal  ring  of  segments,  and"  bisected  by  a  deep  longitudinal  median 
impression,  their  surface  covered  with  microscopic  points.  Thoracic  segments  before  mid- 
length  with  a  transverse  row  of  stiff,  dark  brown  hairs  grouped  in  tufts  or  pencils;  these 
hairs  occurring  on  abdominal  segments  also,  but  less  prominent  here  and  occupying  posterior 
region  of  segments;  pencils  more  numerous  on  ventral  and  pleural  regions,  much  scantier 
on  dorsal  surface  except  on  pronotum.  On  sides  of  last  abdominal  segment,  between  spiracles 
and  spiracular  lobes,  several  long  setae  arranged  in  a  transverse  row,  lacking  on  mid-dorsal 
and  ventral  regions.  Spiracular  disk  (Plate  LXII,  321)  reduced  to  two  ventral  lobes,  as  in 
this  division,  these  lobes  slightly  united  basally,  not  very  elongate,  blunt  at  their  tips,  which 
bear  about  six  dark  setae.  The  two  spiracles  large  and  conspicuous,  entirely  exposed  on 
dorsum  of  last  segment,  separated  by  a  distance  a  little  less  than  diameter  of  one;  middle 
piece  of  spiracles  large,  black;  ring  brownish  yellow.  Anal  gills  four,  long  and  slender,  nearly 
hyaline,  divided  into  lobes  by  slight  constrictions;  posterior  pair  of  lobes  a  little  longer  than 
anterior  pair. 

Head  capsule  elongate,  flattened,  massive,  as  in  this  tribe.  Labrum  conspicuous,  exceed- 
ing men  turn  and  hypopharynx,  the  anterior  margin  fringed  with  long  hairs.  Mentum 
(Plate  LXII,  316)  completely  divided,  each  half  with  three  subequal  prominent  teeth  and 
an  additional  much  smaller  lateral  tooth;  middle  tooth  of  each  side  a  little  broader.  Hypo- 
pharynx  (Plate  LXII, 317)  labriform,  anterior  margin  deeply  concave,  roughened.  Antenna 
(Plate  LXII,  318)  short,  the  segment  short-cylindrical,  a  little  enlarged  toward  truncated 
apex;  at  tip  several  papillae,  of  which  two  are  exceedingly  elongate,  very  slender,  about  twice 
length  of  basal  segment.  Mandible  (Plate  LXII,  319)  powerful,  ending  in  an  acute  point; 
ventral  cutting  edge  with  about  four  teeth,  the  basal  one  of  which  is  the  largest,  subtruncate, 
the  next  outer  tooth  a  large,  flattened,  acute  blade,  two  or  three  smaller  flattened  teeth  just 
before  tip;  dorsal  cutting  edge  with  about  two  small  teeth.  Maxilla  '  (Plate  LXII,  310) 
short  and  very  stout,  the  outer  lobe  much  longer  than  the  slightly  smaller  inner  lobe. 

Pupa.- —  (The  following  notes  are  taken  from  the  cast  pupal  skins  of  the  bred  specimens 

Labrum  truncate.     Labial  lobes  ovate.     Maxillary  palpi  short  and  stout,  broadest  ju?t 
beyond  base,  tips  blunt  (Plate  LXII,  322).     Antenna  moderate  in  length,  tapering  gradu 
ally    to  tip.     Pronotal    breathing    horn  (Plate  LXII,  323  and  324)  short,  cylindrical  or 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  903 

slightly  flattened,  with  a  row  of  circular  breathing  pores  along  margin  of  distal  end,  which 
is  slightly  enlarged.     Wing  sheaths  short.     Leg  sheaths  much  longer. 

Pleural  region  of  abdominal  segments  with  circular  areas  which  are  armed  with  abundant, 
short,  straight  or  slightly  arcuated,  rows  of  spicules,  there  being  from  five  to  seven  spicules 
in  each  row;  under  high  magnification  these  appearing  as  flattened,  scalelike  tubercles, 
fringed  with  the  spicules  described  above;  dorsum  of  intermediate  abdominal  segments 
with  a  broad  transverse  band  which  is  rather  densely  set  with  tiny,  sharply  curved  spines; 
on  posterior  segments  these  rows  lacking  or  much  weaker;  dorsal  bands  lying  posterior  to 
level  of  lateral  areas.  Male  cauda  (Plate  LXII,  325)  with  dorsal  plate  rather  small,  each 
half  indistinctly  bilobed  at  tip;  outer  lobe  minutely  tuberculate  and  with  a  small  seta; 
ventral  lobes  large,  blunt  at  tips;  eighth  tergite  on  either  side  with  a  small  tubercle  bearing 
two  setae. 

Nepionotype. —  Ithaca,  New  York,  May  14,  1917. 

Neanotype. —  Ithaca,  May  26,  1917.  Cast  pupal  skin;  larva  placed  in  rearing  May  14 
1917,  emerged  May  26,  male. 

Paratypes. —  Larvae,  with  type  larva.  Pupae,  taken  as  larvae  May  27,  1917,  placed  in 
rearing,  emerged  June  7,  1917.  No.  79  -  1917. 

Genus  Tricyphona  Zetterstedt  (Gr.   three  +  bend) 

1837  Tricyphona  Zett.     Isis  von  Oken,  p.  65. 

1856  Amalopis  Hal.     Ins.  Brit.,  Dipt.,  vol.  3,  p.  15. 

1856  Bophrosia  Rond.     Dipt.  Ital.  Prodr.,  vol.  1,  p.  183. 

1860  Crunobia  Kol.     Wien.  Ent.  Monatschr.,  vol.  4,  p.  391. 

1881  Nasiterna  Wall.     Ent.  Tidskr.,  vol.  2,  p.  179,  191. 

Larva. —  Body  moderately  elongate,  with  ventral  transverse  creeping- welts  on  basal  annuli 
of  abdominal  segments  4  to  7.  Spiracular  lobes  two,  moderately  elongate.  Anal  gills  four, 
divided  into  two  to  four  lobes  by  from  one  to  three  constrictions.  Head  capsule  massive, 
elongate.  Eyespots  distinct.  Mandible  powerful,  ending  in  an  acute  point  and  with  about 
four  lateral  teeth.  Maxilla  stout,  consisting  of  two  distinct  lobes,  the  slender  inner  lobe 
closely  approximated  to  the  larger  outer  lobe.  Antenna  small;  basal  segment  cylindrical 
at  tip,  with  two  exceedingly  long,  hyaline  papillae  (possibly  lacking  in  T.  immaculata). 
Hypopharynx  labriform,  anterior  margin  concave,  roughened.  Men  turn  completely  divided, 
anterior  margin  of  each  half  with  three  large  teeth. 

Pupa.- —  Pronotal  breathing  horns  short  and  stout,  blunt,  angles  rounded.  Abdominal 
segments  with  transverse  bands  of  spicules  on  tergites  and  sternites,  and  circular  areas 
on  pleurites. 

Tricyphona  is  the  largest  genus  of  the  Pediciini,  including  more  than 
fifty  described  species.  Almost  all  of  these  species  are  found  in  the 
Holarctic  region,  but  two  or  three  are  Antipodal.  In  the  eastern  United 
States,  Tricyphona  inconstans  is  the  most  widely  distributed  and  appar- 
ently the  commonest  species.  This  species  and  certain  others  (as  T. 
paludicola  Alex.)  are  characteristic  swamp  inhabitants,  but  other  species 


904  CHARLES  PAUL  ALEXANDER 

occur  along  running  streams  (T.  vernalis  [0.  S.])  or  near  cliffs  ( T.  auripennis 
[O.  S.]). 

The  immature  stages  of  the  known  species  are  usually  spent  in  moist 
earth.  In  Europe,  T.  immaculata  (Meig.)  was  reared  by  Beling  (1879:47) 
from  larvae  taken  beneath  decaying  vegetable  mold  in  the  bed  of  a 
dried-up  woodland  stream.  Other  specimens  were  found  in  old  horse 
manure  in  beech  woods.  De  Meijere  (1916:195-196)  found  the  same 
species  in  decaying  leaves  and  other  vegetable  matter  in  wet  spots  near 
the  banks  of  streams.  T.  schineri  (Kol.)  was  found  on  August  19  in  a 
wet  spot  in  beech  woods,  where  the  larvae  were  associated  with  pupae 
of  Pedicia  rivosa  in  damp  earth  beneath  'debris,  adults  emerging  on 
September  6  and  12  (Beling,  1879:47).  The  only  American  species 
whose  immature  stages  have  been  found  is  T.  inconstans,  described 
herein. 

Tricyphona  immaculata  (Meig.),  the  genotype,  as  described  by  De 
Meijere  in  the  paper  cited  above,  differs  from  T.  inconstans  as  herein 
described  in  the  following  points:  The  antenna  bears  a  very  short  terminal 
papilla  instead  of  the  two  very  long  ones  in  the  local  species.  The  small 
lateral  tooth  on  the  mentum  is  not  mentioned  nor  figured  as  appearing 
in  T.  immaculata.  The  anal  gills  are  short  and  with  but  a  single  con- 
striction. The  pupa  has  the  pronotal  breathing  horns  small,  kidney- 
shaped,  the  outer  margin  rounded.  The  fore  legs  are  a  little  shorter  than 
the  middle  legs,  and  these  in  turn  are  somewhat  shorter  than  the  posterior 
legs.  The  skin  of  the  abdomen  is  very  delicate,  with  crossrows  of  very 
small  spicules,  arranged  in  numerous  irregular  transverse  rows,  at  the 
posterior  margins  of  the  segments,  about  seven  such  crossrows  being 
present. 

Tricyphona  inconstans  (O.  S.) 

1859    Amalopis  inconstans  0.  S.    Proc.  Acad.  Nat.  Sci.  Phila.,  p.  247. 

Tricyphona  inconstans  is  an  abundant  species  thruout  eastern  North 
America.  It  has  been  recorded  also  from  Europe,  but  the  latter  records 
are  almost  certainly  erroneous.  The  writer  found  larvae  of  this  species 
in  rich  mud  at  Larch  Meadows,  near  Ithaca,  New  York,  on  May  15, 
1917,  in  association  with  larvae  of  Rhamphidia  mainensis,  Pseudolimnophila 
luteipennis,  and  P.  inornata.  Adults  emerged  on  May  27,  1917  (No. 
52-1917).  An  account  of  the  association  in  which  these  larvae  occurred 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  905 

is   given  in   connection   with   the   discussion   of  Rhamphidia  mainensis 
(page  831). 

Larva. —  Length,  17-17.5  mm. 
Diameter,  1.3  mm. 

Color  pale  yellowish  white;  anterior  parts  of  body  sometimes  a  more  saturated  yellow. 
Form  moderately  stout,  body  a  little  narrowed  toward  ends;  surface  of  body  almost 
glabrous,  the  vestiture  being  a  microscopic  pale  pubescence  and  scanty  scattered  hairs. 
Ventral  surface  of  abdominal  segments  4  to  7  with  a  prominent  transverse  welt,  which  is 
hollowed  out  medially  so  as  to  appear  as  paired  prolegs,  these  being  unarmed  with  hooks 
or  points.  Spiracular  lobes  short,  divergent,  tapering  gradually  to  the  blunt  tips,  which  are 
provided  with  seven  or  eight  setae.  Spiracles  (Plate  LXIII,  330)  on  a  transverse  oval  eleva- 
tion, small,  rounded,  separated  by  a  distance  greater  than  diameter  of  one;  middle  piece  of 
spiracles  large,  black,  rings  narrow.  Anal  gills  four,  very  long  and  slender,  constricted 
into  about  four  lobes  which  are  successively  narrowed  from  the  base  outward,  the  terminal 
division  very  slender. 

Head  capsule  of  the  usual  elongate,  massive  type  of  this  tribe.  Labrum  large,  projecting 
beyond  hypopharynx,  anterior  margin  with  long  hairs.  Mentum  (Plate  LXIII,  326)  large, 
completely  divided,  each  half  with  three  subequal  narrow  teeth  and  an  additional  reduced 
lateral  tooth;  on  proximal  margin  of  each  half  of  men  turn,  near  base  of  innermost  tooth, 
a  few  tiny  notches  dovetailing  into  those  of  opposite  half.  Hypopharynx  (Plate  LXIII,  327) 
projecting  far  beyond  men  turn,  labriform;  anterior  margin  deeply  concave  and  provided  with 
small  rounded  papillae;  lateral  angles  smooth,  rounded,  subchitinized.  Antenna  (Plate 
LXIII,  328)  small;  basal  segment  cylindrical,  bearing  at  its  tip  two  elongate  papillae  which 
are  about  one-half  longer  than  basal  segment  alone;  besides  these  an  oval  papilla.  Mandible 
(Plate  LXIII,  329)  of  the  usual  pediciine  type,  powerful,  the  apex  running  out  in  a  long, 
curved  point;  ventral  cutting  edge  very  flat,  cut  into  about  five  teeth,  the  two  basal  of  which 
are  very  large;  the  most  basal  of  these  teeth  squarely  truncated,  the  left  mandible  with  an 
additional  small  tooth  on  lower  inner  angle  of  this  blade;  the  next  outer  tooth  rather  acute; 
outermost  teeth  small  and  flattened;  dorsal  cutting  edge,  as  usual  with  this  division,  with 
two  very  small  teeth  located  far  out  on  apical  point;  a  tuft  of  a  few  long  setae  on  back  of 
mandible  near  heel.  Maxilla  (Plate  LXIII,  329)  powerful,  as  in  this  group  of  genera, 
but  not  exserted  from  the  prothoracic  orifice  when  head  is  retracted;  outer  lobe  very 
stout,  feebly  chitinized  basally,  hyaline  at  apex,  which  bears  the  flattened,  disklike  palpus; 
inner  lobe  slenderer,  shorter,  with  a  few  setae  and  small  papillae. 

Pupa.—  (The  following  notes  are  taken  from  the  cast  skins  of  the  reared  specimens.) 

Labrum  with  apex  broadly  triangular.     Labial  lobes  large,  ovate,  tips  narrowed  and 
bluntly  rounded  (Plate  LXIII,  331).     Pronotal  breathing  horns  (Plate  LXIII,  332  and  333) 
very  short,  stout,  roughly  cylindrical,  the  apices  truncated;  in  lateral  outline,  pentagonal, 
with  a  row  of  breathing  pores  around  margin  of  truncate  apex.     Thoracic  dorsum  trans 
versely  roughened  by  short,   irregular  grooves. 

Abdominal  sternites  with  broad  transverse  bands  of  spicules  on  segments  5  to  7,  the  last 
of  these  three  bands  the  weakest;  similar  bands  on  tergites  4  and  5,  these  bands  subequal 
in  size;  pleural  region  with  a  large,  roughly  circular  area  of  short  rows  of  microscopic  spicules, 


906  CHARLES  PAUL  ALEXANDER 

these  areas  near  base  of  segments.  Female  cauda  (Plate  LXIII,  334)  with  tergal  valves 
elongate,  tapering  gradually  to  the  rather  blunt  tips;  sternal  valves  very  small,  blunt  at  tips; 
at  base,  on  dorsal  side  of  eighth  segment,  two  setiferous  tubercles. 

Nepionotype  —  Larch  Meadows,  Ithaca,  New  York,  May  15,  1917.     No.  52-1917. 
Neanotype. —  Ithaca,  May  27,  1917,  a  cast  pupal  skin. 

Group  Dicranotae 

Genus  Dicranota  Zetterstedt  (Gr.  a  fork) 

1838     Dicranota  Zett.     Ins.  Lapponica,  Dipt.,  p.  851,  no.  164. 

Dicranota  is  a  small  genus  of  crane-flies  (about  fifteen  described  species) 
occurring  thruout  the  Holarctic  region.  Four  species  are  found  in  eastern 
North  America.  None  of  the  American  species  have  been  reared.  In 
Europe,  the  life  history  and  morphology  of  Dicranota  bimaculata  (Schum.) 
has  been  discussed  in  detail  by  Miall  (1893)  and  by  Wesenberg-Lund 
(1915:342-343).  Larvae  were  found  by  Miall  in  numbers  in  the  muddy 
banks  of  small  streams  and  ponds,  where  they  bury  themselves  in  the 
mud  and  gravel.  They  creep  about  with  ease  and  rapidity  between  the- 
sancl  and  gravel,  and  are  able  to  swim  well  by  a  looping  movement.  Their 
food  consists  largely  of  small  worms,  Tubifex  rivulorum  Lam.,  which 
abound  in  these  haunts.  The  pupal  stage  is  passed  in  damp  earth.  The 
larva  is  stated  to  be  about  18  millimeters  in  length,'  but  this  seems  to  be 
a  maximum  figure.  The  body  is  dirty  white  in  color  and  is  covered  with 
fine,  appressed  hairs.  Abdominal  segments  3  to  7  bear  paired  retractile 
pseudopods,  which  are  circled  with  three  rows  of  chitinized  hooks  gradually 
decreasing  in  size  from  the  tips  inward.  The  anal  gills,  four  in  number, 
are  distinctly  segmented.  The  head  capsule  is  elongate  and  massive, 
as  in  the  tribe.  The  mentum  is  completely  divided,  its  anterior  margin 
having  the  usual  six  teeth.  The  mandible  is  of  the  usual  pediciine  type, 
with  acute  teeth  on  the  ventral  cutting  edge  and  a  brush  of  hairs  near 
the  prosthecal  region.  (Plate  LXIV.) 

The  pupa  is  small,  only  about  10  millimeters  in  length,  and  has  the 
pronotal  breathing  horns  expanded  and  flattened  at  the  tips.  The  dorsal 
surface  of  the  abdomen  is  provided  with  roughened  plates  armed  with 
rather  strong  and  dense  spines,  there  being  one  such  plate  on  the  third 
segment,  two  on  the  fourth  to  sixth  segments,  and  one  on  the  seventh 
segment. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  907 

Genus  Rhaphidolabis  Osten  Sacken  (Gr.  needle  +  forceps) 

1869    Rhaphidolabis  0.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  281-2~7. 
1911     Claduroides  Brun.     Rec.  Indian  Mus.,  vol.  6,  p.  288. 

Rhaphidolabis  is  a  small  genus,  including  about  fifteen  described  species 
ranging  thruout  the  North  Temperate  Zone.  The  larvae  strongly  resemble 
those  of  Dicranota,  and  the  two  genera  are  undoubtedly  very  closely 
related. 

Rhaphidolabis  tenuipes  O.  S. 

1869    Rhaphidolabis  tenuipes  0.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  287. 

Needham  (1908 a: 212-214)  found  adult  flies  of  the  species  Rhaphidolabis 
tenuipes  in  great  numbers  in  tent  traps  set  over  Beaver  Meadow  Brook 
in  the  Adirondacks  in  July  and  August,  1907.  Larvae  that  almost  cer- 
tainly belong  here  were  found  among  the  rounded  stones  in  the  creek 
bottom. 

Larva. —  Length  excluding  caudal  lobes,  8-9  mm. 
Length  of  caudal  lobes,  1  mm. 

Abdominal  segments  3  to  7  with  prominent  fleshy  prolongations  on  ventral  surface,  these 
being  retractile,  unpaired,  and  widely  separated  on  mid-ventral  line,  and  bearing  at  tip  a 
circlet  of  outcurved  booklets,  with  series  of  smaller  booklets  beyond.  Spiracular  disk  with 
the  two  ventral  lobes  long  and  obtuse  at  tips.  (Further  details  are  given  by  Needham.) 

Rhaphidolabis  cayuga  Alex,  (supposition) 

1916    Rhaphidolabis  cayuga  Alex.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  543-544. 

Larvae  which  the  writer  refers  to  this  species  were  found  in  Needham's 
Glen,  Ithaca,  New  York,  on  March  30,  1916.  Later  in  the  season,  in 
April  and  May,  adult  flies  of  the  species  are  very  numerous  in  this  glen, 
and  these  account  for  the  specific  reference. 

Larva.- —  Agreeing  very  closely  with  descriptions  of  bbth  Dicranota  and  Rhaphidolabis 
tenuipes.  Spiracles  large,  lying  in  a  distinct  longitudinal  groove,  by  the  closing  of  which  they 
are  capable  of  being  entirely  hidden.  Antenna  long  and  slender,  tapering  to  apex,  which 
bears  numerous  short  papillae.  Mandible  with  third  lateral  tooth  of  ventral  cutting 
row  very  long  and  acute,  much  exceeding  second  tooth;  basal  tooth  shaped  like  a  prun- 
ing knife,  with  the  cutting  edge  sinuate;  at  prosthecal  region  of  mandible,  a  small  tuft  of 
about  seven  or  eight  long  setae.  Maxilla  having  the  two  lobes  firmly  united  except  on 
apical  quarter. 


908  CHARLES  PAUL  ALEXANDER 

Tribe  Eriopterini 

The  tribe  Eriopterini  comprises  a  vast  assemblage  of  usually  small  crane- 
flies  whose  geographical  range  is  coextensive  with  that  of  the  family. 

The  larvae,  so  far  as  known,  are  herbivorous.  They  show  a  remarkable 
uniformity  in  the  structure  of  the  head  capsule.  In  the  generalized 
members  such  as  Molophilus  and  some  Erioptera,  the  ventral  bars  of  the 
head  capsule  are  toothed  at  their  anterior  ends  and  form  mental  plates 
which  are  apparently  homologous  with  those  of  the  Pediciini.  In  Chionea 
an  apparently  similar  condition  exists,  to  judge  from  Brauer's  figures. 
In  the  majority  of  species,  however,  the  ventral  bars  of  the  capsule  are 
not  enlarged  nor  toothed  anteriorly  and  do  not  function  as  the  mental 
plates.  The  hypopharynx  is  preserved  as  a  hemispherical  cushion  which 
is  densely  provided  with  setae.  The  mandibles  are  small  and  are  blunt 
at  their  tips;  the  teeth  of  the  cutting  edge  are  usually  three  in  number 
and  blunt,  but  in  some  species  (Ormosia,  Gonomyia)  they  are  longer 
and  more  prominent.  A  distinct  prosthecal  lobe  or  hook  is  usually 
developed,  and  near  the  base  of  the  mandible  is  a  slitlike  opening  bearing 
a  fringe  of  long,  yellow  setae.  The  maxillae  are  rather  large,  hairy  lobes. 
The  labrum  and  epipharynx  is  long  and  narrow,  and  densely  hairy.  The 
antennae  are  remarkably  uniform  thruout  the  group,  consisting  of  a  stout 
cylindrical  basal  segment  with  a  slightly  smaller  apical  papilla  of  an 
elongate-oval  shape.  In  the  Elephantomyaria  the  condition  is  somewhat 
similar,  the  mandibles  being  very  small,  and  the  esophageal  region  being 
conspicuously  grooved  with  parallel  lines  and  ridges.  The  spiracular 
disk  is  obliquely  truncated  and  indistinctly  lobed  in  Chionea  and  Teucho- 
labis;  surrounded  by  four  lobes  in  Elephantomyia;  squarely  truncated 
and  surrounded  by  four  (in  Rhabdomastix)  or  usually  five  lobes.  In 
the  undetermined  Eriopterine  No.  1,  the  five  lobes  are  spatulate  flattened 
blades  with  the  margins  hooked.  Anal  gills  are  usually  present  and 
variously  developed  in  the  different  genera.  The  pupa  is  rather  slender, 
in  the  Elephantomyaria  with  the  rostral  sheath  very  long  and  the  palpi 
strongly  recurved.  The  head  usually  bears  a  small  setiferous  crest  which 
is  rarely  lacking.  The  pronotal  breathing  horns  vary  considerably  in 
form  and  relative  size,  being  flattened  into  fans  in  some  Gonomyia,  small 
and  trumpet-shaped  in  other  Gonomyia  and  in  Gnophomyia,  elongate 
in  most  species.  The  mesonotum  is  usually  armed  at  the  crest  with  spines, 
hooks,  or  setiferous  tubercles.  The  leg  sheaths  are  very  short  in  Gno- 


THE  CRANE-FLIES  OF  NEW  YORK — PART  II  909 

phomyia  but  are  longer  in  other  genera,  and  the  middle  tarsi  are  usually 
shorter  than  the  other  legs.  The  abdominal  segments  are  subdivided 
into  two  rings,  the  posterior  ring  with  a  transverse  row  of  spines  and 
setae  before  the  margin.  The  lateral  spiracles  are  small,  protuberant, 
and,  in  some  cases  at  least,  apparently  functional.  The  dorsum  of  the 
eighth  abdominal  segment  is  provided  with  four  or  five  lobes,  which  are 
in  some  cases  (as  in  some  Ormosia)  spinous  at  the  tips. 

The  writer  has  subdivided  the  tribe  Eriopterini  into  two  divisions: 
the  Elephantomyaria,  with  Elephantomyia  and  presumably  Toxorhina 
and  Ceratocheilus,  and  the  Eriopteraria,  including  the  other  Nearctic 
genera  as  known.  It  is  probable  that  Cladura  and  its  relatives  will  require 
a  division  when  their  immature  stages  are  better  known. 

The  genera  of  the  tribe  Eriopterini  may  be  separated  by  the  following 

keys: 

Larvae 

1.  Spiracular  disk  surrounded  by  four  lobes 2 

Spiracular  disk  not  as  aoove 3 

2.  Ventral  lobes  with  a  single  powerful  seta;  coloration  a  saturated  golden  yellow;  lives 

under  bark Elephantomyia  O.  S.  (p.  952) 

Ventral  lobes  fringed  with  short  setae;  coloration  pale  yellow;  lives  in  earth. 

Rhabdomastix  Sk.  (p.  942) 

3.  Spiracular  disk  obliquely  truncated,  appearing  indistinctly  trilobed  or  without  lobes. .  .4 
Spiracular  disk  squarely  truncated,  surrounded  by  five  lobes 5 

4.  Mandibles  with  eight  teeth;  mental  plates  with  seven  teeth;  lives  in  earth. 

Chionea  Dalman  (p.  950) 
Mandibles  with  less  than  eight  teeth;  mental  plates  not  toothed;  lives  under  bark. 

Teucholabis  O.  S.  (p.  945) 

5.  Spiracular  disk  ending  in  five  flattened  black  plates  which  are  finely  toothed  along  their 

margins Genus  incertus,  Eriopterine  No.  1  (p.  956) 

Spiracular  disk  not  as  above 6 

6.  Ventral  plates  of  head  capsule  expanded  and  toothed  at  anterior  ends 7 

Ventral  plates  not  toothed  as  above . .-° 

7.  Ventral  plates  with  four  teeth;  coloration  yellow;  spiracular  disk  large,  very  heavily 

marked  with  black Molophilus  Curt.  (p.  911) 

Ventral  plates  with  five  to  eight  teeth;  coloration  green;  spiracular  disk  very  reduced, 
almost  unmarked Erioptera  (chlorophylla  0.  S.)  (p.  918) 

8.  Marks  of  all  the  lobes  solidly  black Trimicra  O.  S.  (p.  932) 

Some  Ormosia  Rond.  (p.  922) 
Some  of  the  marks  more  or  less  split  by  a  pale  line -9 

9.  The  three  dorsal  lobes  solidly  dark,  the  ventral  pair  split  by  a  pale  line 10 

All  paired  lobes  split  by  a  pale  line J1 

10.  Coloration  saturated  yellow;  anal  gills  bluntly  rounded;  lives  under  bark. 

Gnophomyia  O.  S.  (p.  934) 

Coloration  pale  yellow;  anal  gills  elongate;  lives  in  earth. 

Helobia  St.  Farg.  et  Serv.  (p.  928) 

11.  Lobes  surrounding  disk  very  stout,  blunt;  marks  of  lateral  lobes  surrounding  sPiracle? 

and  often  suffusing  disk Gonomyia  Meig.,  subgenus  Leiponeura  Skuse  (p.  VM) 

Lobes  surrounding  disk  longer;  marks  of  lobes  not  so  extensive ** 


910  CHABLES  PAUL  ALEXANDER 

12.  All  the  lobes  with  two  lines 13 

Dorsal  lobe  solidly  dark Some  Ormosia  Rond.  (p.  922) 

13.  No  dark  marks  on  disk  between  spiracles Some  Ormosia  Rond.  (p.  922) 

Dark  marks  between  spiracles 14 

14.  Spiracular  disk  very  small;  two  spots  on  disk.  .  .  Erioptera  (megophthalma  Alex.)  (p.  915) 
Spiracular  disk  large;  four  or  six  spots  on  disk Helobia  St.  Farg.  et  Serv.  (p.  928) 

Pupae 

1.  Rostral  sheath  very  elongated;  antennal  sheaths  lying  across  face  of  eye. 

Elephantomyia  O.  S.  (p.  952) 
Rostral  sheath  not  elongated;  antennal  sheaths  lying  behind  eyes 2 

2.  Leg  sheaths  very  short,  barely  exceeding  wings;  crest  of  mesonotum  smooth  or  nearly  so. 

Gnophomyia  O.  S.  (p.  934) 

Leg  sheaths  longer,  ending  about  one  segment  beyond  end  of  wings;  crest  of  mesonotum 
with  spines  or  tubercles 3 

3.  Mesonotum  at  crest  with  numerous  sharp  spines;  hind  tarsi  the  longest,  middle  tarsi  the 

shortest,  fore  tarsi  intermediate  in  length Helobia  St.  Farg.  et  Serv.  (p.  928) 

Mesonotum  and  tarsal  arrangement  not  as  above 4 

4.  Mesonotum  at  crest  with  six  to  eight  tubercles  provided  with  numerous  setae;  breathing 

horns  fanlike  or  trumpet-like Gonomyia  Meig.  (p.  938) 

Mesonotum  and  breathing  horns  not  as  above 5 

5.  Mesonotum  at  crest  with  four  sharp  spines  or  two  plates  produced  into  spines 6 

Mesonotum  at  crest  without  distinct  prominent  spines 8 

6.  Mesonotum  at  crest  with  four  spines 7 

Mesonotum  at  crest  with  a  plate  on  either  side  split  at  tip  into  three  or  four  teeth. 

Ormosia  (nubila  [O.  S.j)  (p.  923) 

7.  Crest  of  mesonotum  with  two  large  spines;  pronotal  breathing  horns  small,  closely  applied 

to  thorax;  lives  under  bark Teucholabis  O.  S.  (p.  945) 

Crest  of  mesonotum  with  four  large  spines;  pronotal  breathing  horns  long,  slender,  curved; 
lives  in  earth Molophilus  Curt.  (p.  911) 

8.  Abdominal  pleurites  with  a  transve/se  subterminal  row  of  from  eight  to  twelve  spines  or 

stout  setae;  dorsum  of  segment  8  with  a  pentagon  of  five  lobes. 

Ormosia  Rond.  (p.  922) 

Abdominal  pleurites  without  such  a  row  of  spines  or  setae;  dorsum  of  segment  8  with  four 
or  fewer  lobes Erioptera  Meig.  (p.  914) 

The  most  important  literature  on  the  tribe  Eriopterini  is  as  follows: 

Molophilus  bijttatus Larva,  general Keilin,  1913:4.     (Hypodermal 

glands.) 

Molr,philus  obscurus General Beling,  1879  56. 

Molophilus  ochraceus Larva,  pupa Beling,  1886: 193-194. 

Ei  ioptera  maculata Pupa Beling,  1879 : 49. 

Erioptera  flavescens Larva Beling,  1879: 50. 

Erioptera  lutea Larva,  pupa Beling,  1886: 192-193. 

Erioptera  sordida General Beling,  1879 : 56. 

Cheilotrichia  imbuta General De  Meijere,  1920: 76. 

Lip.wlhrix  icterica Larva,  pupa Beling,  1886: 192. 

Ormosia  haemorrhoidalis Pupa Beling,  1879 : 48^19. 

Ormosia  haemorrhoidalis Larva,  pupa Beling,  1886: 191-192. 

Ormosia  lineata General Beling,  1886: 202. 

Ormosia  nodulosa General Beling,  1886: 202. 

Ormosia  nodulosa General Cameron,  1917: 65. 

Ormosia  varia Larva,  pupa De  Meijere,  1916:201-204. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  911 

Helobix  hybrida Larva,  pupa Beling,  1879: 50-51. 

Helobia  hybrida Larva Hart,  1898  [1895]:  199-200. 

Helobia  hybrida Larva,  pupa Malloch,  1915-17  b:  229-230. 

Trimicra  pilipes Larva,  general Gerbig,  1913: 161-163. 

Gnophomyia  rufa Larva,  pupa Hudson,  1920: 32-33. 

Gnophomyia  tripudians General Gamkrelidze,  1913,  a  and  b. 

Gnophomyia  tripudians General.  , Keilin,  1913:3.     (Hypodermal 

glands.) 

Gnophomyia  tripudians Larva Keilin,  1913:4.     (Hypodermal 

glands.) 

Gnophomyia  tripudians General Edwards,  1919  b. 

Gnophomyia  tristissima Larva,  pupa,  general. . .  Malloch,  1915-17 b: 230-231. 

Gonomyia  tenella General Beling,  1879: 56. 

Rhnbdomastix  schistacea Larva,  pupa Beling,  1886: 195. 

Trentepohlia  bromeliadicola Larva,  pupa,  general. . .  Picado,  1913:356-357. 

Trentepohlia  leucoxena General Alexander,  1915  b. 

Trentepohlia  pennipes Larva,  pupa,  general . . .  De  Meijere,  1911:50-51. 

Teucholabis  complexa General Johnson,  1900. 

Chionea  araneoides Larva,  general Brauer,  Egger,  and  Frauenfeld, 

1854. 

Subtribe  Eriopteraria 

Genus  Molophilus  Curtis  (derivation  obscure) 
1833    Molophilus  Curt.     Brit.  Ent.,  p.  444. 

Larva. —  Form  long  and  slender.  Spiracular.  disk  squarely  truncated  and  surrounded  by 
five  subequal  lobes;  paired  lobes  of  disk  heavily  lined  with  black;  a  black  mark  running  proxi- 
mad  from  spiracles;  dorsal  lobe  with  an  oval  black  area.  Head  capsule  long  and  narrow; 
ventral  rods  flattened;  anterior  ends  expanded,  four-toothed,  to  form  mental  plates. 
Labrum  narrow,  densely  hairy.  Mandible  blunt  at  apex;  ventral  cutting  edge  with  about 
fo'ir  teeth;  a  single  subapical  dorsal  tooth.  Maxilla  large  and  blunt.  Antenna  large;  apical 
papilla  elongate-oval,  sculptured. 

Pupa. —  Cephalic  crest  setiferous.  Pronotal  breathing  horns  long  and  slender,  sinuously 
curved.  Mesonotum  declivitous,  at  crest  with  four  powerful  teeth.  Wing  sheaths  ending 
at  about  midlength  of  second  abdominal  segment.  Leg  sheaths  ending  at  about  midlength 
of  fourth  abdominal  segment;  tarsi  of  middle  legs  the  shortest.  Abdomen  with  spiracles 
on  segments  2  to  7;  dorsum  of  segment  8  with  four  blunt  lobes. 

Molophilus  is  a  well-defined  genus  (including  more  than  fifty  species) 
of  small  and  usually  dull-colored  flies,  which  are  found  practically  thruout 
the  world. 

The  adult  flies  frequent  shaded  situations  and  may  be  swept  from 
rank  vegetation  in  such  places.  The  swarming  habits  of  this  group  are 
discussed  later  under  the  specific  accounts. 

The  immature  stages  of  the  various  species  are  spent  in  wet  earth. 
In  Europe,  Molophilus  obscurus  (Meig.)  (Beling,  1879:56)  and  M.  ochraceus 
(Meig.)  (Beling,  1886:193-194)  have  been  found  in  wet  earth  near  running 


912  CHARLES  PAUL  ALEXANDER 

water  in  shaded  woods.  The  pupal  duration  of  the  latter  species  is  not 
more  than  ten  days.  Keilin  (1913:4)  notes  the  presence  of  hypodermal 
glands  in  larvae  of  M.  bifilatus  Verr.,  but  does  not  mention  the  larval 
habitat.  Of  the  American  species,  M.  hirtipennis  has  been  reared  from 
similar  situations. 

Molophilus  hirtipennis  (O.  S.) 

1859     Erioptera  hirtipennis  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  228. 

The  little  dark-colored  crane-flies  of  the  species  Molophilus  hirtipennis 
are  common  on  vegetation  in  shaded  woods  in  spring  and  early  summer, 
or  they  may  be  found  in  small  dancing  swarms  in  similar  situations. 

The  larvae  are  exceedingly  abundant  in  wet  organic  mud  or  in  cool,  rich 
woods  in  the  neighborhood  of  streams  or  springs.  The  writer  has  reared 
the  species  very  frequently  from  specimens  found  in  Needham's  Glen 
and  on  Bool's  hillside,  Ithaca,  New  York,  in  April  and  May,  1914  to 
1917.  The  pupal  period  is  probably  about  a  wrck,  but  in  all  the  rearings 
of  the  writer  this  could  not  be  ascertained  closer  than  ten  days. 

Larva. —  Length,  9-10  mm. 

Diameter,  0.4-0.5  mm. 

Coloration  light  yellow. 

Form  long  and  narrow;  body  terete,  noticeably  constricted  before  spiracular  disk  (Plate 
XLV,  344).  Integument  covered  with  a  delicate  appressed  pubescence  and  a  few  transverse 
rows  of  very  short,  erect  setae.  Spiracular  disk  (Plate  LXV,  351)  squarely  truncated,  sur- 
rounded by  five  subequal  lobes;  ventral  lobes  on  inner  face  with  two  heavy  black  parallel 
lines,  separated  by  a  capillary  yellow  line;  lateral  lobes  with  a  similar  double  line  running 
inward  far  beyond  spiracle;  these  double  lines  not  connected  at  distal  end;  dorsal  lobe  with 
a  single  oval  black  mark  which  is  less  intense  outwardly;  a  black  mark  beginning  at  spiracle 
running  proximad  toward  center  of  disk;  lobes  with  a  few  short  hairs  at  tips.  (There  is  a 
little  variation  in  the  degree  of  intensity,  but  the  general  pattern  is  as  described  above.) 
Anal  gills  four,  short  and  blunt. 

Head  capsule  (Plate  LXV,  345)  long,  narrow,  consisting  of  six  chitinized  rods,  the  ventral 
rods  broad  and  flat,  at  the  anterior  end  expanded  to  form  the  mental  plates  (Plate  LXV,  346), 
each  rod  contributing  four  teeth,  of  which  the  outermost  is  bluntly  rounded,  the  middle  pair 
the  largest  and  subequal.  Dorsal  bars  two  on  either  side,  one  slender,  at  their  anterior 
ends  articulating  with  a  transverse  chitinized  rod;  the  various  bars  connected  by  a  thin 
membranous  tissue.  Labrum  and  epipharynx  elongate,  narrow,  occupying  the  space  between 
mandible  and  antenna  on  either  side,  the  ventral  face  with  abundant  long  hairs,  on  sides 
margined  with  numerous  long,  incurved,  flattened  setae.  Mental  plates  as  described  above; 
behind  them  the  hypopharynx  (Plate  LXV,  347),  consisting  of  a  semicircular  cushion  provided 
with  dense,  short  setae.  Antennae  (Plate  LXV,  348)  rather  closely  approximated  on  dorsum, 
prominent,  each  1-segmented  but  bearing  a  long  apical  papilla;  basal  segment  moderately 
elongated,  cylindrical,  the  apex  obliquely  truncated,  the  papilla  hyaline,  gradually  narrowed 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  913 

toward  tip,  shaped  somewhat  like  an  ear  of  corn,  the  surface  delicately  sculptured;  besides 
this  papilla,  two  or  three  much  smaller  cylindrical  ones.  Mandible  (Plate  LXV,  349)  slender, 
ending  in  a  blunt  rounded  lobe;  ventral  cutting  edge  with  four  blunt  teeth,  the  second  from 
the  base  very  tiny  and  connected  with  the  third  from  the  base  (in  some  specimens  the  teeth 
are  very  blunt  and  rounded,  so  that  the  cutting  edge  appears  crenulated  or  wavy);  a  single 
tooth  on  dorsal  cutting  edge,  immediately  behind  apex;  prosthecal  region  of  mandible  with 
a  longitudinal  slitlike  opening  filled  with  a  dense  row  of  long  setae.  Maxilla  (Plate  LXV, 
350)  large  and  blunt,  the  outer  lobe  pale,  roughly  triangular,  covered  with  numerous  short 
hairs,  and  with  a  few  small  sensory  papillae  near  apex,  surrounding  palpus;  inner  lobe  a 
little  shorter,  densely  clothed  and  fringed  with  abundant  long  yellow  hairs. 

Pupa.—  Length,  6.2-7.3  mm. 

Width,  d.-s.,  0.7-0.8  mm. 
Depth,  d.-v.,  0.8-1  mm. 

Head,  thorax,  and  appendages  brown,  when  fully  matured  almost  black;  breathing  horns 
yellowish;  abdomen  pale  yellowish  white. 

Cephalic  crest  of  moderate  size,  consisting  of  two  conical  lobes  directed  forward  and  bearing 
a  stout  seta  on  anterior  face;  immediately  in  front  of  these,  a  smaller  transverse  crest  lying 
between  antennal  bases.  Front  rather  broad.  Labrum  triangular;  lobes  of  labium  triangular, 
divergent;  maxillary  palpi  elongate,  moderately  stout,  narrowed  at  tips  and  extending  beyond 
joints  of  fore  legs.  Thorax  prominent,  carinate  medially  before  declivity.  Breathing 
horns  long  and  slender,  sinuously  curved,  apices  directed  forward.  Declivity  of  mesonotum 
(Plate  LXVI,  352)  precipitous,  at  the  crest  armed  with  four  powerful  teeth,  median  pair  the 
largest,  somewhat  divergent,  separated  by  median  line;  a  few  tiny  setae  on  mesonotum,  includ- 
ing four  in  a  transverse  row  at  about  the  level  of  wing  root.  Wing  sheaths  ending  opposite 
apex  of  second  abdominal  segment.  (In  fully  matured  pupae  the  characteristic  venation  of 
this  genus  shows  on  the  wing  pads.)  Leg  sheaths  rather  short,  ending  about  opposite  mid- 
length  of  fourth  abdominal  segment;  hind  legs  slightly  the  longest,  fore  legs  a  little  shorter, 
middle  legs  conspicuously  shorter,  ending  about  opposite  apex  of  fourth  tarsal  segment  of 
fare  legs. 

Abdominal  segments  with  two  very  narrow  basal  rings  and  a  much  broader  posterior 
ring.  Abdominal  spiracles  distinct  on  segments  2  to  7.  Setae  as  follows:  on  pleura,  one 
immediately  caudad  of  spiracle,  a  second  caudad  and  somewhat  dorsad  of  spiracle  opposite 
pasterior  ring,  another  opposite  anterior  ring.  Tergum  with  two  setae  on  sides  of  posterior 
ring.  Segment  8  with  four  blunt  lobes  on  dorsum.  Male  cauda  (Plate  LXVI,  353  and  354) 
with  ventral  lobe  large,  bluntly  rounded  at  tips;  dorsal  lobes  ending  in  two  acute,  chitinized 
points  which  are  widely  separated  and  directed  dorsad  and  slightly  caudad,  on  outer  face 
with  a  small  seta;  near  base  of  cauda  on  dorsum,  surface  tumid  and  bearing  a  small  seta  on 
either  side.  Female  cauda  (Plate  LXVI,  355  and  356)  with  sternal  acidothecae  much  shorter 
than  the  very  long  tergal  valves,  these  latter,  just  before  apex,  with  an  acute  spine  which  is 
directed  dorsad  and  with  a  seta  on  side.  (When  the  pupa  is  nearly  ready  to  transform 
to  the  adult,  the  long,  coarse  bristles  covering  the  body  of  the  adult,  and  the  chitinized 
genitalia,  show  thru  the  pupal  integument.) 

Nepionotype  —  Ithaca,  New  York,  May  11,  1917.     No.  33-1917. 

Neanotype. —  Ithaca,  June  1,  1917. 

Paratypes  —  About  one  hundred  larvae  and  pupae  from  type  locality. 

8 


914  CHARLES  PAUL  ALEXANDER 

Molophilus  ursinus  (0.  S.) 

1859     Erioptera  ursina  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  228. 

Molophilus  ursinus  is  probably  the  smallest  crane-fly  in  North  America. 
The  following  notes  on  the  swarming  were  made  along  Power  House 
Creek,  Gloversville,  New  York,  on  June  27,  1915: 

This  species  occurred  in  small  dancing  swarms  over  the  little  lateral  streamlets  that  poured 
down  the  steep  slope,  some  of  the  swarms  including  scores  of  individuals.  When  not  swarm- 
ing, they  rested  flat  on  the  upper  surfaces  of  leaves,  their  wings  folded  over  the  abdomen. 
Crane-flies  associated  with  this  species  today  included  the  following:  BittacomorpheUa 
jonesi,  Limnophila  toxoneura,  L.  areolata,  L.  (Lasiomastix)  tenuicornis,  Liogma  nodicornis, 
Dolichopeza  americana,  Tipula  collaris,  T.  nobilis,  T.  iroquois,  T.  hermannia,  T.  macrolabis, 
T.  submaculata,  T.  fuliginosa,  Nephrotoma  macrocera,  N.  tennis,  Longurio  testaceus,  and  others. 

Genus  Erioptera  Meigen  (Gr.  wool  +  wing) 

1800  Polymeda  Meig.     Nouv.  Class.  Mouch.,  p.  14  (nomen  nudum). 

1803  Erioptera  Meig.     Illiger's  Mag.,  vol.  2,  p.  262. 

1818  Polyraphla  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  1,  p.  88. 

1854  Octavia  Bigot.     Ann.  Soc.  Ent.  France,  p.  474. 

1856  Chemalida  Rond.     Dipt.  Ital.  Prodr.,  vol.  1,  p.  180. 

1856  Limnaea  Rond.     Dipt.  Ital.  Prodr.,  vol.  1,  p.  181. 

1856  Ilisia  Rond.     Dipt.  Ital.  Prodr.,  vol.  1,  p.  182.    r 

1861  Limnoica  Rond.     Dipt.  Ital.  Prodr.,  Corrigenda,  vol.  4,  p.  11. 

1863  Trichosticha  Schin.     Wien.  Ent.  Monatschr.,  vol.  7,"  p.  221. 

1864  Platytoma  Lioy.    Atti  dell'  Institut  Veneto,  ser.  3,  vol.  10,  p.  42. 

Larva. —  Form  elongate,  in  some  cases  very  slender,  terete.  Spiracular  disk  tending  to 
be  reduced,  in  some  species  (as  E.  chlorophylla)  very  small.  Anal  gills  blunt.  Head  capsule 
slender,  consisting  of  six  bars,  four  dorsal  and  two  ventral;  ventral  bars  in  some  species 
(as  E.  chlorophy lla) ,  at  least,  with  five  to  eight  teeth  at  their  expanded  anterior  end,  forming 
the  mental  plates.  Labrum  narrow,  densely  hairy.  Mandible  small,  bluntly  toothed. 
Maxilla  blunt,  hairy.  Hypopharynx  cushion-like. 

Pupa.- —  Cephalic  crest  consisting  of  blunt  or  acute  (in  E.  chlorophylla  and  E.  vespertina) 
lobes.  Pronotal  breathing  horns  elongate,  the  length  many  times  the  diameter,  usually 
straight  and  slightly  divergent  (E.  megopkthalma,  E.  septemtrionis) ,  or  acutely  pointed  and 
directed  forward  (E.  chlorophylla).  Mesonotal  declivity  along  crest  with  very  numerous 
setiferous  tubercles  or  (in  the  European  E.  lutea)  with  four  prominent  teeth.  Leg  sheaths 
moderately  long,  reaching  fourth  abdominal  segment;  middle  tarsi  conspicuously  shorter 
than  the  others.  Subapical  armature  of  abdominal  segments  strong  on  sternites  and  tergites 
but  lacking  on  pleurites.  Spiracles  distinct,  tubular.  Lobes  on  dorsum  of  eighth  abdominal 
segment  not  forming  a  distinct  pentagon. 

Erioptera  is  a  large  genus  of  small  flies,  including  more  than  one  hundred 
described  species,  chiefly  from  the  temperate  regions.  In  the  case  of 
all  species  of  which  the  immature  stages  are  known,  these  stages  are  spent 
in  wet  earth.  In  Europe,  Erioptera  (Acyphona)  maculata  (Beling,  1879: 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  915 

49)  was  found  under  leaves  in  damp  earth  in  woods.  Pupae  taken  on 
June  8,  1874,  emerged  as  adults  on  the  13th.  The  species  was  found 
also  in  the  sandy  mud  of  a  small  brook  bed.  E.  (E.)  flavescens  (Beling, 
1879:50)  was  found  in  sandy,  damp  earth,  E.  (E.)  sordida  (Beling,  1879: 
56,  mention  only)  in  wet,  sandy  earth  along  brooks,  and  E.  (E.)  lutea 
(Beling,  1886:192-193)  in  woods,  in  damp  hollows,  and  along  the  margins 
of  brooks,  usually  under  leaf  mold.  Edwards  (1919  a)  has  recorded  a 
species  of  Erioptera  as  being  associated  with  the  larvae  of  the  mosquito 
Taeniorhynchus  at  the  roots  of  water  grass  (Glyceria  fluitans)  in 
England. 

In  America,  E.  chlorophylla,  E.  vespertina,  E.  caloptera,  and  other 
species  inhabit  wet  mud  in  open  swamps;  E.  septemtrionis  and  E.  megoph- 
thalma, rich  organic  mud  in  cool,  shady  woods;  E.  armata,  E.  near 
knabi,  and  other  species,  the  soil  along  the  sandy  banks  of  streams.  E. 
(Acyphona)  graphica  was  considered  by  Hart  (1898  [1895]:  197)  to  be 
semiaquatic. 

The  writer  has  before  him  the  larvae  of  only  two  species  of  the  genus, 
E.  chlorophylla  and  E.  megophthalma.  These  are  readily  separated  by 
their  color,  E.  chlorophylla  being  pale  green,  E.  megophthalma  pale 
yellow  and  more  elongate.  The  pupae  of  the  Nearctic  species  of  the 
genus  may  be  separated  by  the  following  key: 

1.  Breathing  horns  and  cephalic  crest  strongly  pointed  and  curved  forward  at  their  tips; 

coloration  light  green E.  chlorophylla  O.  S.  (p.  918) 

Breathing  horns  and  cephalic  crest  not  as  above;  coloration  not  green 2 

2.  Breathing  horns  elongate-cylindrical  to  slightly  flattened,  almost  straight  but  slightly 

divergent,  diameter  subequal  for  entire  length 3 

Breathing  horns  not  as  above,  more  or  less  curved,  or  else  blunt  at  their  tips 4 

3.  Cephalic  crest  sharply  pointed;  eyes  large E.  megophthalma  Alex.  (p.  915) 

Cephalic  crest  not  sharply  pointed;  eyes  small E.  septemtrionis  0.  S.  (p.  920) 

4.  Breathing  horns  almost  straight,  enlarged  distally,  tips  blunt. 

E.  sp.  (near  knabi  Alex.)  (p.  922) 
Breathing  horns  curved,  narrowed  toward  tips 5 

5.  Breathing  horns  widely  separated  at  base,  bent  strongly  proximad,  contiguous  medially; 

lobes  of  cephalic  crest  acute,  vertical;  eyes  large.  .  , E.  vespertina  O.  S.  (p.  919) 

Breathing  horns  curved  gently  forward;  cephalic  crest  with  lobes  directed  laterad,  diver- 
gent; eyes  of  moderate  size E.  armata  O.  S.  (p.  921) 

Erioptera  megophthalma  Alex. 

1918     Erioptera  megophthalma  Alex.     Can.  Ent.,  vol.  50,  p.  60-61. 
The    adult    flies    of  Erioptera  megophthalma    are    not    uncommon   in 
cool,    shaded,   and  boggy   woods    during  the    months   of   early   spring. 


916  CHARLES  PAUL  ALEXANDER 

They  may  be  swept  from  rich  vegetation  in  company  with  such  char- 
acteristic species  as  Limnophila  brevifurca,  L.  fuscovaria,  L.  subtenuicornis, 
Adelphomyia  minuta,  Rhaphidolabina  flaveola,  Molophilus  hirtipennis, 
Erioptera  venusta,  E.  stigmatica,  Gonomyia  florens,  G.  subcinerea,  and 
Tipula  oropezoides. 

The  elongate  larvae  of  this  fly  were  very  common  in  rich  organic  mud 
taken  from  Bool's  hillside,  Ithaca,  New  York  (as  discussed  under  the 
account  of  Bittacomorphella  jonesi,  page  780),  where  they  were  associated 
with  a  crane-fly  fauna  characteristic  of  such  situations.  The  larvae,  in 
life,  are  pale  yellowish,  with  the  food  contents,  of  a  chalky  white  color, 
showing  thru  the  integument.  The  head  capsule  and  the  spiracular  disk 
are  very  small ;  the  inner  face  of  each  lobe  of  the  latter  is  very  narrowly 
lined  with  black.  The  species  was  reared  many  times  during  late  May 
and  early  June,  1917,  the  length  of  the  pupal  existence  indoors  being  seven 
or  eight  days. 

Larva. —  Length,  10.4-11.6  ram. 
Diameter,  0.7-0.75  mm. 

Coloration  very  pale  yellow;  contents  of  alimentary  canal  chalky  white. 

Form  terete,  elongated,  body  tapering  gradually  to  the  posterior  end,  just  beyond  gills 
(Plate  LXVII,  353)  suddenly  constricted;  last  segment  elongate-cylindrical,  tapering  gradu- 
ally to  the  very  small  spiracular  disk.  Body  covered  with  a  short,  appressed  pubes- 
cence, on  last  segment  this  pubescence  coarser  and  more  erect,  with  a  few  elongate  hairs 
interspersed;  lateral  parts  of  body  at  caudal  margins  of  segments  with  short  transverse 
lines  of  small,  erect  setae;  a  few  other  similar  rows  at  about  midlength  of  certain  of  the 
segments.  Spiracular  disk  (Plate  LXVII,  357)  very  small,  tending  to  be  eliminated  by 
reduction;  lobes  short  and  blunt,  dorso-median  lobe  the  smallest;  ventral  lobes  with  two 
short  brown  lines,  not  connected  distally,  the  proximal  line  a  little  longer  than  the  lateral 
line  of  each  lobe;  the  pale  space  between  these  lines  a  little  less  than  diameter  of  one; 
lateral  lobes  with  two  similar  divergent  lines,  the  dorsal  one  attaining  inner  level  of 
spiracles;  dorsal  lobe  with  two  small,  indistinct,  brown  lines;  on  disk  between  spiracles  two 
small  round  spots  which  do  not  touch  spiracles;  lobes  fringed  with  short  hairs  near  tip, 
and  capable  of  close  approximation  so  that  disk  is  often  entirely  closed.  Spiracles  large, 
nearly  circular. 

Head  capsule  small,  very  long  and  slender,  greatly  dissected,  the  three  bars  of  either  side 
long  and  delicate;  dorsal  bars  at  their  articulation  joined  with  a  short  longitudinal  bar  near 
whose  anterior  end  the  antennae  are  inserted;  ventral  bars  of  capsule  not  conspicuously 
expanded  at  their  anterior  end,  and  apparently  not  toothed  as  in  other  species  of  this  genus 
and  in  Molophilus.  Labrum  and  epiphirynx  long  and  narrow,  lying  between  antennal 
bases;  epipharyngeal  region  densely  clothed  with  short  setae  at  tip  and  with  two  parallel 
brushes  on  ventral  face.  Mentum  apparently  not  formed  as  in  E.  chlorophylla,  a  slightly 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  917 

arched  transverse  chitinized  bar.  Hypopharynx  about  as  in  Molophilus.  Antennae  rather 
closely  approximated,  directed  cephalad;  basal  segment  moderately  elongated,  cylindrical; 
apical  papilla  relatively  small,  elongate-oval.  Mandible  very  small,  with  blunt  teeth; 
apical  point  short,  blunt;  ventral  row  of  teeth  about  three  in  number,  often  very  blunt. 
Maxilla  as  in  Molophilus,  but  outer  lobe  with  the  vestiture  of  hairs  rather  longer. 

Pupa.—  Length,  7.2-8.2  mm. 

Width,  d.-s.,  0.8-0.9  mm. 
Depth,  d.-v.,  1-1.1  mm. 

Head  light  brown;  thorax  anterior  to  declivity  conspicuously  darker  brown;  remainder 
of  body  light  yellowish  brown;  breathing  horns  light  yellow.  (In  fully  colored  individuals 
the  head  and  the  thorax  with  their  appendages  become  much  darker,  almost  black,  but  the 
breathing  horns  retain  their  conspicuous  yellow  color.) 

Head  short,  face  tumid.  Eyes  of  male  very  large,  widely  separated  by  front;  eyes  of 
female  smaller.  Cephalic  crest  consisting  of  two  prominent  lobes;  viewed  from  side,  these 
lobes  sharply  pointed  and  directed  slightly  forward,  with  a  seta  on  outer  ventral  face  before 
tips;  viewed  from  front,  lobes  rounded,  ending  in  acute  tips,  separated  by  a  deep,  rather 
narrow,  V-shaped  notch.  Front  between  eyes  narrowed  toward  labrum,  which  is  rather 
sharply  pointed.  Labial  sheaths  small,  the  lobes  contiguous  with  their  apices  truncated, 
the  lateral  angles  obtuse  or  produced  into  a  tiny  lobe.  Sheaths  of  maxillary  palpi  short 
and  stout,  tapering  gradually  to  tips.  Antennal  sheaths  moderately  elongated,  angulated 
at  segments,  ending  just  beyond  base  of  wing.  Pronotal  breathing  horns  stout,  expanded 
at  base,  almost  straight  and  only  slightly  divergent,  somewhat  compressed,  transversely 
wrinkled  basally;  a  small  setiferous  tubercle  in  front  of  base  of  breathing  hern.  Mesonotum 
precipitous,  at  crest  (Plate  LXVII,  359)  on  either  side  of  median  line  with  abundant  tiny 
setiferous  tubercles  bearing  long,  pale  hairs,  these  tubercles  continued  back  along  shoulder. 
Lateral  margin  of  thorax  with  two  small  setae.  Wing  sheaths  ending  before  tip  of  second 
abdominal  segment.  Leg  sheaths  (Plate  LXVII,  360)  short,  attaining  base  of  fourth 
abdominal  segment;  tarsi  of  hind  legs  the  longest,  those  of  middle  pair  the  shortest;  fore  legs 
with  femora  and  tibiae  very  short. 

Abdominal  segments  (Plate  LXVII,  361)  divided  into  two  annuli  by  a  constriction  near 
midlength,  the  anterior  ring  very  indistinctly  subdivided  further  into  two  lesser  annulets;  on 
segments  4  to  7,  before  caudal  margin  of  posterior  ring  en  both  dorsum  and  sternum,  a  trans- 
verse row  of  small,  conspicuous,  blackened,  setiferous  tubercles,  which  are  more  distant 
from  one  another  near  ends  of  rows;  on  basal  abdominal  segments  these  tubercles  less  evident 
but  still  present.  Pleura  with  small  but  probably  noa-functional  spiracles,  which  are  very 
indistinct  in  young  pupae  but  are  more  evident  in  fully  colored  individuals;  these  spiracles 
located  near  base  of  posterior  ring.  Setae  on  abdomen  as  follows:  on  sternal  segments, 
one  seta  just  caudad  of  end  of  row  of  spicules,  a  second  at  lateral  end  of  this  row,  inter- 
mixed with  spicules,  two  on  posterior  ring  on  a  level  with  spicules;  on  tergal  segments,  a 
strong  seta  on  a  line  with  spiracles,  another  seta  below  end  of  row  of  spicules;  on  pleura,  one 
seta  just  ventrad  of  spiracles,  and  two  post-spiracular  and  one  ante-spiracular  setae.  Male 
cauda  (Plate  LXVII,  362  and  363)  with  ventral  lobes  obliquely  truncated,  blunt  at  tips, 
projecting  beyond  level  of  subacute  dorsal  lobes;  dorsal  lobes  slender,  slightly  divergent 
apically,  blackened  before  tips  and  with  two  setae  at  tips  on  outer  face;  at  base  of  cauda, 


918  CHARLES  PAUL  ALEXANDER 

on  dorsal  face  of  eighth  segment,  a  close  quadrangle  of  four  rounded  tubercles,  placed  on 
a  slight  elevation.  Female  cauda  with  tergal  acidothecae  elongate,  subacute  at  tips;  sternal 
valves  short,  blunt;  quadrangle  of  tubercles  on  dorsum  of  eighth  segment  more  distinct 
than  in  the  male. 

Nepionotype. —  Ithaca,  New  York,  May  14,  1917. 
Neanotype. —  Ithaca,  June  5,  1917. 

Paratypes. —  Larvae  and  pupae  in  large  numbers  from  type  locality,  May  14  to  June  5, 
1917. 

Erioptera  chlorophylla  O.  S. 

1859     Erioptera  chlorophylla  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  226. 

Erioptera  chlorophylla  is  conspicuous  by  its  pale  green  color  in  the 
larval,  pupal,  and  adult  stages.  It  is  a  common  and  widely  distributed 
species  thruout  eastern  North  America.  Several  larvae  were  found  in 
organic  mud  at  Orono,  Maine,  on  June  13,  1913,  one  of  which  pupated 
on  the  21st.  When  the  insect  is  dropped  into  boiling  water,  the  green 
color  immediately  disappears.  The  associates  of  this  species  are  discussed 
under  the  account  of  Ptychoptera  rufocincta  (page  775). 

Larva. —  Length,  9-10  mm. 

Diameter,  0.7-0.75  mm. 

Color  uniformly  pale  green,  fading  to  a  pale  yellow  after  death. 

Form  moderately  elongated,  last  segment  of  body  elongate,  gradually  narrowed  to  bluntly 
rounded  apex.  Body  clothed  with  numerous  appressed  hairs.  Spiracular  disk  somewhat 
as  in  E.  megophthalma  but  even  more  reduced,  disk  usually  entirely  closed,  lateral  lobes 
on  either  side  capable  of  close  approximation,  tracheae  before  the  opening  into  spiracles 
very  large. 

Head  capsule  (Plate  LXVIII,  364)  of  the  Molophilus  type,  but  longer  and  slenderer;  the 
two  dorsal  bars  of  each  side  very  delicate,  the  ventral  bars  broader  and  flattened.  Mental 
plates  (Plate  LXVIII,  365  and  366)  slender;  anterior  end  of  each  ventral  bar  widely 
expanded  and  provided  with  several  teeth;  these  teeth  varying  in  number,  in  some  speci- 
mens there  being  only  five,  in  others  eight,  teeth  to  each  plate;  in  the  latter  case  the  third 
from  either  side  is  larger,  with  two  smaller  teeth  between.  Hypopharynx  about  as  in 
Molophilus.  Antenna  (Plate  LXVIII,  367)  large,  basal  segment  stout,  cylindrical;  apical 
papilla  elongate-oval,  with  apex  bluntly  rounded  and  surf  ace  weakly  sculptured;  laterad  of 
this  papilla  a  tiny  cylindrical  hyaline  peg.  Mandible  (Plate  LXVIII,  368)  rather  large; 
cutting  edge  with  about  four  slender  teeth,  the  second  from  base  the  smallest;  dorsal  face 
of  mandible  with  a  blunt  subapical  tooth  and  an  oblique  comb  of  about  six  stout  setae  or 
chitinized  teeth.  Maxilla  similar  to  that  of  Molophilus,  but  the  hairy  vestfture  longer  and 
coarser. 

Pupa.—  Length,  8.8-9  mm. 

Width,  d.-s.,  1.2  mm. 
Depth,  d.-v.,  1.2  mm. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  919 

Breathing  horns  reddish  brown;  thoracic  dorsum  green,  with  a  brownish  tinge; 
.abdomen  uniformly  pale  green,  posterior  half  of  each  dorsal  segment  a  little  darker. 

Cephalic  crest  small  and  compact,  consisting  of  two  prominent  but  closely  approximated 
lobes  which  are  separated  by  a  deep  U-shaped  notch,  the  tips  acute  and  directed  forward; 
on  outer  face  before  apex  a  short  seta.  Labrum  broadly  obtuse  at  tip.  Labial  lobes  roughly 
triangular,  divergent.  Maxillary  palpi  short  and  stout,  narrowed  toward  tip.  Antennal 
sheaths  with  a  slender  tubercle  at  base  above  eye. 

Pronotal  breathing  horns  broad  at  base,  narrowed  to  the  acute  tip  which  is  directed  almost 
ventrad;  viewed  from  above,  horns  very  broad  basally  and  with  a  dorsal  carina;  ventral 
side  at  base  transversely  wrinkled;  a  small  setiferous  lobe  just  in  front  of  breathing  horns, 
directed  laterad.  Mesonotum  behind  breathing  horns  with  a  high  compressed  carina,  on 
either  side  of  this  produced  into  a  lobe  directed  cephalad  and  laterad;  mesonotum  moderately 
declivitous  (Plate  LXIX,  369) ,  at  crest  with  numerous  setiferous  tubercles  which  are  fewer 
in  number  and  more  widely  separated  along  shoulder.  Leg  sheaths  reaching  to  about  middle 
of  fourth  abdominal  segment;  hind  legs  a  little  longer  than  fore  legs;  middle  legs  very  short, 
ending  opposite  base  of  last  segment  of  fore  legs. 

Abdominal  segments  divided  into  two  narrow  basal  rings  and  a  broad  posterior  ring; 
on  pleura  a  distinct  spiracle,  opposite  posterior  annulus  and  nearer  dorsal  margin;  posterior 
annulus,  before  caudal  margin,  with  a  dorsal  and  a  ventral  row  of  long,  stout  setae.  Setae 
on  abdomen  as  follows:  on  pleura,  a  seta  opposite  second  basal  ring,  a  second  ventrad  of 
spiracle,  and  two  setae  caudad  of  spiracle,  the  posterior  one  a  little  more  dorsal  in  position; 
on  tergites,  two  stout  setae  lying  transversely  on  the  margin  opposite  spiracle,  a  third  seta 
at  end  of  terminal  rows  of  bristles;  on  sternites,  a  group  of  two  transverse  setae  on  posterior 
ring,  slightly  below  level  of  spiracle  and  rather  widely  separated  by  the  broad  midventral 
area.  Female  cauda  (Plate  LXIX,  370)  with  dorsal  acidothecae  short,  distinctly  upturned, 
and  ending  in  a  small,  subacute  tip;  before  apex  with  two  very  short  setae;  a  short  blunt 
tubercle  near  base  of  valves;  ventral  lobes  short,  their  tips  very  blunt;  dorsum  of  segment  8 
with  two  blunt  median  tubercles,  one  immediately  behind  the  other. 

(Described  from  larvae  taken  in  the  Basin  Swamp,  Orono,  Maine,  June  13,  1913;  one  pupa 
with  the  larva,  July  5,  1913.) 

Erioptera  vespertina  O.  S. 

1859  Erioptera  vespertina  O.  S.  Proc.  Acad.  Nat.  Sci.  Phila.,  p.  226. 
Erioptera  vespertina  is  a  characteristic  inhabitant  of  open  swamps  and 
wet  meadows.  It  has  not  been  reared,  but  a  pupa  found  in  organic  mud 
in  the  Basin  Swamp,  Orono,  Maine,  on  June  24,  1913,  undoubtedly  belongs 
to  this  species.  The  associates  are  discussed  under  the  account  of  Bittaco- 
morpha  davipes  (page  785). 

Pupa. —  Length,  8.5  mm. 

Depth,  d.-v.,  1.1  mm. 

Breathing  horns  reddish  brown;  thoracic  dorsum  reddish  brown,  with  an  interrupted 
whitish  line  running  down  posterior  half  of  mesonotum;  sheaths  of  wings  and  legs  pale 


920  CHARLES  PAUL  ALEXANDER 

brown;  abdomen  whitish  yellow,  posterior  half  of  each  sternite  and  tergite  dark  brown, 
basal  half  with  two  narrow  transverse  lines  of  same  color. 

Lobes  of  cephalic  crest  widely  separated,  subtriangular,  acutely  pointed  at  tips.  Pronotal 
breathing  horns  cylindrical,  transversely  wrinkled,  tapering  gradually  to  tips.  (In  the  only 
specimen  at  hand,  the  horns  are  widely  separated  at  the  base  but  soon  bend  proximad  and 
thence  outward  and  ventrad,  so  that  the  two  horns  are  closely  approximated  or  almost 
contiguous  on  their  distal  parti.)  Behind  breathing  horns,  mesonotum  with  conspicuous 
divergent  lobes  such  as  are  described  for  E.  chlorophylla;  mesonotum  at  crest  with  abundant 
black  setiferous  tubercles,  which  are  fewer  in  number,  smaller,  and  more  scattered  along 
shoulder,  interrupted  at  mid-dorsal  line.  Hind  legs  a  little  longer  than  fore  legs,  which, 
in  turn,  are  a  little  longer  than  middle  legs. 

Abdominal  segments  with  subterminal  armature  of  posterior  ring  more  spinous  than  in 
E.  chlorophylla.  Distribution  of  setae  about  as  in  E.  chlorophylla. 

(Described  from  a  pupa  taken  at  Orono,  Maine,  June  24,  1913.) 

Erioptera  septemtrionis  O.  S. 

1859    Erioptera  septemtrionis  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  226. 

Erioptera  septemtrionis  is  a  widely  distributed  species  thruout  the 
northeastern  United  States  and  Canada.  The  larvae  are  not  uncommon 
in  rich  organic  mud  in  cool,  shaded  woods.  The  writer  found  them 
commonly  in  the  Standpipe  Woods,  Orono,  Maine,  in  July  and  August, 
1913.  A  larva  placed  in  rearing  on  July  3  transformed  to  an  adult  female 
on  the  16th.  Other  larvae  placed  in  rearing  on  July  15  transformed  to 
adult  males  on  the  25th.  This  limits  the  pupal  duration  to  not  more  than 
ten  days,  but  it  is  probably  much  less,  presumably  about  one  week. 

Pupa.—  Length  of  cast  skin,  7  mm. 

Cephalic  crest  low  and  flat.  Antennal  sheaths  moderately  elongated,  individual  segments 
showing  clearly  thru  sheaths.  Pronotal  breathing  horns  elongate,  cylindrical,  almost  straight 
but  slightly  diverging,  transversely  wrinkled,  paler  at  tips,  with  a  row  of  small  breathing 
pores  along  apical  margin;  a  few  small  setiferous  tubercles  before  base  of  breathing  horns. 
Declivity  of  mesonotum  rather  steep,  at  crest  with  numerous  small  tubercles  and  abundant 
pale  yellow  hairs  which  are  less  numerous  along  shoulder.  Leg  sheaths  with  middle  pair 
conspicuously  the  shortest,  as  in  the  genus. 

Abdominal  segments  with  subterminal  armature  of  posterior  ring  consisting  of  stout, 
pale  setae;  arrangement  of  these  setae  about  as  in  E.  chlorophylla.  Lateral  spiracles  distinct. 
Male  cauda  (Plate  LXIX,  371  and  372)  with  the  ventral  lobes  blunt  at  tips,  obliquely 
truncated,  separated  by  an  acute  V-shaped  notch;  dorsal  lobes  separated  by  a  U-shaped 
notch,  each  lobe  terminating  in  a  small  tip,  with  two  tiny  setae  on  lateral  face  before 
apex;  dorsum  of  eighth  segment  with  four  prominent,  pale,  fleshy  lobes  which  are  closely 
approximated,  the  anterior  pair  directed  laterad  and  a  little  more  distant  from  each  other 
than  the  posterior  pair,  which  are  directed  more  dorsad. 

Neanotype.—  Orono,  Maine,  July  25,  1913.     No.  105-1913. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  921 

(Subgenus  Hoplolabis  Osten  Sacken) 

1869    Hoplolabis  0.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  160. 

The  subgenus  Hoplolabis  includes  but  three  known  species  —  the  type 
of  the  group,  Erioptera  (Hoplolabis)  armata,  discussed  below;  E.  (H.) 
bipartita  O.  S.,  of  western  North  America;  and  E.  (H.)  asiatica  Alex., 
of  Japan. 

Erioptera  (Hoplolabis)  armata  O.  S. 

1859     Erioptera  armata  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  227. 

Erioptera  armata  is  a  rather  common  fly  thruout  the  northeastern 
United  States.  A  larva  was  found  in  the  sand  along  the  banks  of  Fall 
Creek,  Ithaca,  New  York,  on  May  16,  1917.  This  larva  was  of  the 
typical  eriopterine  form,  being  elongate,  terete,  and  with  the  spiracular 
disk  surrounded  by  five  subequal  lobes.  It  was  placed  in  rearing  and 
transformed  to  an  adult  female  on  May  31.  This  larva  was  found  associ- 
ated with  numerous  hexatomine  larvae,  such  as  Eriocera  spinosa,  E. 
longicornis,  and  E.  cinerea. 

The  following  description  is  from  the  cast  pupal  skin. 

Length,  about  7  mm. 

Cephalic  crest  consisting  of  two  moderately  large,  slightly  divergent  lobes  which  are 
acutely  pointed  at  tips;  lobes  directed  strongly  outward  and  bearing  a  seta  on  outer  face. 
Labrum  acutely  pointed.  Labial  lobes  large,  divergent,  almost  straight  across  caudal  margin. 
Sheaths  of  maxillary  palpi  stout,  rather  pointed  at  tips.  Pronotal  breathing  horns  broad 
at  base,  tapering  to  slender  apices,  bases  conspicuously  wrinkled;  a  small  setiferous 
tubercle  before  base  of  each  breathing  horn.  Mesonotum  at  crest  (Plate  LXIX,  373) 
with  numerous  short,  chitinized  points.  Lateral  angle  of  thorax  with  two  setae;  a  strong 
seta  above  wing  axil.  Leg  sheaths  with  middle  tarsi  ending  conspicuously  before  tarsi  of 
other  legs. 

Each  abdominal  segment  before  posterior  margin  with  a  transverse  row  of  slender  black 
spicules  or  short  bristles  on  tergum  and  sternum,  and  smaller  areas  on  pleura.  Lateral 
abdominal  spiracles  distinct.  Setae  on  pleura  just  ventrad  of  spiracle,  and  on  tergum  just 
above  transverse  row  of  spicules.  Female  cauda  with  tergal  valves  elongated,  gently 
upcurved,  unarmed;  sternal  valves  shorter,  blunt;  at  base  of  cauda,  on  dorsum  of  eighth  seg- 
ment, four  small  darkened  tubercles  which  are  produced  into  slender  tips. 

Neanotype.*—  Cast  pupal  skin,  Ithaca,  New  York,  May  31,  1917. 

(Subgenus  Mesocyphona  Osten  Sacken) 

1869    Mesocyphona  O.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  161. 
Mesocyphona  is  one  of  the  larger  subgenera  of  Erioptera,  reaching  its 
maximum  of  specific  development  in  the  Tropics  of  the  New  World. 


922  CHARLES  PAUL  ALEXANDER 

The  immature  stages  of  Erioptera  (Mesocy phono)  cdloptera  (Say)  and 
E.  (M.)  parva  O.  S.  are  spent  in  wet  mud  along  the  banks  of  streams 
and  other  bodies  of  water.  The  species  discussed  below  as  Erioptera 
(Mesocyphona)  species  (near  knabi),  was  reared  from  the  sandy  margins 
of  a  small  prairie  stream  in  Kansas. 

Erioptera  (Mesocyphona)  species  (near  knabi  Alex.) 

Adult  flies  of  a  small  species  of  Mesocyphona  which  is  close  to  E.  (M.) 
knabi  Alex.,  of  Mexico,  were  not  uncommon  along  Buckner  Creek,  a  small 
prairie  stream  flowing  thru  Jetmore,  Kansas.  These  adults,  especially 
the  females,  were  photophilous,  appearing  in  considerable  numbers  around 
lanterns  which  were  hung  in  tents  pitched  along  the  banks  of  this  stream'. 
A  single  pupa  found  in  the  muddy  sand  along  the  bank  of  the  creek  on 
July  20,  1917,  emerged  as  an  adult  on  the  22d.  The  following  general 
characters  of  the  species  may  be  noted: 

Pupa. —  Labrum  small,  apex  rather  sharp.  Labial  lobes  squarely  truncated,  with  lateral 
angles  subacute.  Sheaths  of  maxillary  palpi  slender.  Pronotal  breathing  horns  moderately 
long,  cylindrical,  curved  slightly  forward,  enlarged  outwardly,  blunt  at  tips.  Mesonotum 
at  crest  rather  tumid  and  with  a  few  long  hairs  inserted  on  stout  black  tubercles.  Leg  sheaths 
with  middle  tarsi  the  shortest,  hind  tarsi  the  longest. 

Neanotype.—  Jetmore,  Hodgeman  County,  Kansas,  July  22,  1917. 

Genus  Ormosia  Rondani  (Gr.  chain) 

1856    Ormosia  Rond.     Dipt,  Ttal.  Prodr.,  vol.  1,  p.  180. 

1860    Rhypholophus  Kol.     Wien.  Ent.  Monatschr.,  vol.  4,  p.  393. 

1863     Dasyptera  Schin.     Wien.  Ent.  Monatschr.,  vol.  7,  p.  221. 

Larva.' — Form  terete,  moderately  elongated.  Spiracular  disk  squarely  truncated,  sur- 
rounded by  five  subequal  lobes  which  are  lined  with  double  marks  of  brown.  Anal  gills 
blunt.  Head  capsule  slender,  very  dissected,  of  six  narrow  bars,  four  dorsal  and  two  ventral, 
the  ventral  bars  broader.  Labrum  narrow,  epipharynx  hairy.  Mandible  with  teeth 
moderately  elongated.  Antenna  of  the  Molophilus  type.  Mentum  without  chitinized 
teeth. 

Pupa.- —  Cephalic  crest  setiferous.  Pronotal  breathing  horns  rather  short  and  stout, 
more  or  less  flattened  and  with  a  row  of  tubercles  along  posterior  margin.  Mesonotum 
declivitous,  at  crest  with  a  flattened,  toothed,  chitinized  plate  on  either  side  (in  0.  nubila] 
or  with  abundant  setiferous  tubercles.  Wing  sheaths  ending  opposite  or  just  beyond  tip 
of  second  abdominal  segment.  Leg  sheaths  varying  in  length  with  the  different  species, 
middle  tarsi  the  shortest.  Abdominal  segments  with  a  subterminal  transverse  row  of  spines 
or  setae,  these  occurring  on  pleura  (as  small  groups  of  eight  to  twelve)  as  well  as  on  tergites 
and  sternites.  Lateral  spiracles  distinct,  on  segments  2  to  7;  dorsum  of  segment  8  with 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  923 

a  pentagon  of  five  lobes,  these  being  unarmed  (0.  innocens,  0.  meigenii)  or  spinous-tipped 
(0.  nubila,  0.  nigripila). 

Ormosia  is  a  large  and  rather  difficult  genus  (including  more  than 
seventy-five  species)  of  small  crane-flies  whrch  are  characteristic  of  sub- 
arctic and  temperate  regions  and  apparently  rare  or  lacking  in  the  Tropics. 

The  adult  flies  occur  in  small  dancing  swarms,  usually  in  cool,  shaded 
situations  in  or  near  woods  or  along  brooks.  They  are  most  numerous 
in  early  spring  and  in  late  summer  or  early  autumn,  many  of  the  species 
being  apparently  double-brooded. 

The  immature  stages  are  spent  in  moist  organic  mud  near  water.  In 
Europe,  Beling  records  Ormosia  haemorrhoidalis  (Zett.)  (Beling,  1879: 
48-49,  and  1886:191-192),  0.  lineata  (Meig.)  (Beling,  1886:202),  and 
0.  nodulosa  (Macq.)  (Beling,  1886:202),  as  being  found  in  wet  earth  in 
woods,  usually  beneath  leaf  mold.  Cameron  (1917 : 65)  likewise  records  the 
last-named  species  as  living  in  mud.  0.  varia  (Meig.)  was  found  by  De 
Meijere  (1916:201-204)  among  decaying  leaves  in  a  wet  spot  near  a  ditch. 

Of  the  American  species,  the  writer  has  bred  Ormosia  innocens,  0. 
nubila,  0.  meigenii,  and  0.  nigripila  from  larvae  or  pupae  in  entirely 
similar  situations  to  those  given  above  for  the  European  species. 

Tho  larvae  of  only  two  of  these  species,  0.  nubila  and  0.  meigenii,  are 
available  to  the  writer  at  this  time.  0.  nubila  is  a  large  brown  species; 

0.  meigenii  is  much  slenderer,  and  is  light  yellow  in  color. 

The  pupae  of  the  known  Nearctic  species  may  be  distinguished  as 
follows: 

1.  Dorsum  of  eighth  abdominal  segment  with  a  pentagon  of  five  lobes  which  are  spinous 

at  their  tips 2 

Dorsum  of  eighth  abdominal  segment  with  a  pentagon  of  five  fleshy  lobes 3 

2.  Crest  of  mesonotum  with  a  flattened  chitinized  plate  on  either  side  of  median  line. 

0.  nubila  (0.  S.)  (p.  923) 

Crest  of  mesonotum  tumid,  with  abundant  coarse,  yellowish  setae  on  either  side  of  median 
line 0.  nigripila  (0.  S.)  (p.  927) 

3.  Abdomen  with  large,  rectangular,  dusky  areas  on  posterior  annuli  of  segments  2  to  8, 

giving  abdomen  a  banded  appearance;  pleural  setae  rather  numerous. 

0.  innocens  (O.  S.)  (p.  925) 
Abdomen  without  such  dusky  areas:  pleural  setae  few  in  number. 

0.  meigenii  (O.  S.)  (p.  928) 

Ormosia  nubila  (0.  S.) 

1859     Erioptera  nubila  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  227. 

Ormosia  nubila  is  probably  double-brooded,  since  the  flies  are  on  the 
wing  in  the  spring  and  again  in  the  fall.  The  immature  stages  live  in 


924  CHARLES  PAUL  ALEXANDER 

organic  mud  that  is  usually  covered  over  with  a  layer  of  leaf  mold.  On 
March  27,  1914,  the  writer  found  four  large  brown  eriopterine  larvae  in 
rich  earth  from  Needham's  Glen,  Ithaca,  New  York.  An  adult  female 
of  the  present  species  emerged  on  April  9.  This  makes  the  maximum 
pupal  existence  less  than  two  weeks,  bu.t  it  is  undoubtedly  very  much 
less  than  this. 

Larva. —  Length,  11.5-11.8  mm. 
Diameter,  1.1-1.2  mm. 

Color  a  deep  reddish  or  cinnamon  brown;  incisures  of  segments  paler. 

Form  rather  stout,  body  terete.  Skin  covered  with  a  short,  appressed  pubescence. 
Spiracular  disk  (Plate  LXX,  375)  squarely  truncated,  surrounded  by  five  subequal  lobes; 
ventral  lobes  with  two  parallel  dark  brown  lines  which  are  narrowly  united  distally,  the  pale 
stripe  between  rather  broad,  distinct,  especially  near  center  of  disk;  lateral  lobes  with  two 
parallel  dark  brown  stripes  which  are  indistinctly  connected  distally,  the  space  between 
dusky  with  numerous  brown  spots;  dorsal  lobe  with  an  elongate-oval  mark  inclosing  a  linear 
yellow  center;  lobes  fringed  with  long  hairs  which  are  longest  at  tips,  shorter  toward  base, 
and  narrowly  interrupted  between  lobes;  disk  between  spiracles  unmarked.  Spiracles  large; 
middle  piece  and  extreme  outer  margins  of  ring  blackish;  spiracles  separated  by  a  distance 
about  equal  to  one  and  one-half  times  diameter  of  one. 

Head  capsule  about  as  in  Molophilus,  but  ventral  bars  of  capsule  not  toothed  to  form 
the  characteristic  mental  plate  of  that  genus.  Labrum  and  epipharynx  about  as  in  Molo- 
philus. Hypopharynx  broad,  flattened,  provided  with  numerous  transverse  rows  of  short 
setae.  Antenna  short;  basal  segment  stout,  cylindrical;  apical  papilla  rather  small, 
elongate-oval.  Mandible  (Plate  LXX,  374)  ending  in  a  rather  long  apical  point,  with  about 
four  long,  flattened  teeth  along  ventral  cutting  edge,  the  second  from  base  very  small;  a  short, 
recurved  hook  at  prosthecal  region  and  a  dense  tuft  of  long  yellow  hairs  in  prosthecal  slit;  a 
dorsal  appendage  at  heel  of  mandible.  Maxilla  about  as  in  Molophilus. 

Pupa. —  Length,  7.5  mm. 

Width,  d.-s.,  1.2  mm. 
Depth,  d.-v.,  1.4  mm. 

Head,  thorax,  and  appendages  pale  yellow;  breathing  horns  yellow;  chitinized  plates  on 
mesonotum  dark  brown;  abdomen  brown.  (In  mature  pupae,  the  sheaths  of  the  appendages 
are  probably  darker.) 

Cephalic  crest  (Plate  LXXI,  381)  low;  lateral  angles  produced  into  conical,  erect, 
spinous  tubercles,  each  bearing  a  stout  seta  on  outer  ventral  face.  Antennal  sheaths  very 
angulated,  almost  serrate.  Opposite  each  segment  of  antenna  on  basal  half  of  organ,  a 
conspicuous  blackened  tubercle,  those  at  base  larger  and  more  conspicuous,  the  lateral  one 
directed  outward,  above  it  a  second  tubercle  directed  cephalad  and  simulating  a  crest. 
Antenna  extending  to  just  beyond  base  of  wings.  Front  broad;  a  blackish  area  on  either 
side  near  inner  margin  of  eye,  probably  indicating  point  of  attachment  of  tentorium.  Labrum 
triangular,  apex  subacute.  Labial  lobes  triangular,  tips  blunt.  Sheaths  of  maxillary  palpi 
rather  long,  narrowed  to  the  slender  tip.  Pronotal  breathing  horns  short  and  slender,  some- 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  925 

what  flattened,  pale  yellow;  a  stout  seta  just  before  base  of  each  breathing  horn.  Mesonotum 
precipitous;  at  crest  (Plate  LXX1,  380)  on  either  side  of  median  line,  a  heavily  chitinized 
flattened  plate  projecting  dorsad;  this  plate,  at  its  tip,  forking  into  two  lesser  teeth,  the 
lateral  one  of  these  still  further  subdivided  into  two  still  smaller  teeth  (in  one  specimen 
the  left  plate  is  divided  dichotomously  into  four  teeth,  the  inner  primary  tooth  being  further 
subdivided) ;  on  shoulder  laterad  of  these  plates  a  slightly  swollen  plate  which  is  parallelly 
grooved;  two  spines  on  lateral  angle  of  thorax  above  base  of  wing;  mesonotum  with  a  few 
setae  behind  crest,  as  follows:  an  anterior  solitary  seta  en  either  side  of  median  line  and  close 
to  it,  a  solitary  stout  seta  above  wing  axil,  two  groups  of  paired  setae,  one  just  dorsad  of  base 
of  wing  and  the  other  midway  between  this  group  and  the  anterior  seta  first  described.  Wing 
sheaths  rather  short,  ending  opposite  base  of  third  abdominal  segment.  Leg  sheaths  rather 
long  for  this  genus,  ending  opposite  midlength  of  fifth  abdominal  segment;  fore  tarsi  a  little 
shorter  than  hind  tarsi,  middle  tarsi  very  short,  ending  opposite  or  just  beyond  end  of  third 
tarsal  segment  of  fore  legs. 

Abdominal  segments  near  posterior  margin  with  transverse  rows  of  short  black  spineg 
which  are  much  smaller  and  more  widely  separated  near  ends  of  row;  at  intervals  alona 
row  a  few  elongate  setae;  on  pleura  a  similar  area  of  eight  to  eleven  spines  with  a  single  sets 
in  row;  tubular  lateral  spiracles  on  segments  2  to  7.  Chaetotaxy  as  follows:  on  pleura,  just 
above  dorsal  end  of  row,  one  seta,  another  solitary  seta  cephalad  of  spiracle  on  anterior 
ring;  on  tergites,  a  solitary  seta  on  posterior  ring  about  opposite  spiracle;  on  sternites,  two 
small  setae  on  either  side  of  median  line  of  posterior  ring.  Female  cauda  (Plate  LXXI,  382) 
with  tergal  valves  long  and  slender,  almost  straight  but  slightly  upcurved  near  tips;  on 
dorsal  lateral  margin,  just  before  tip,  a  prominent  tooth  directed  laterad  and  slightly  caudad; 
about  midlength  of  valves  a  somewhat  similar  blunt  tubercle  on  either  side  near  dorsal  margin; 
at  base  of  eighth  tergite  a  pentagon  of  five  chitinized  lobes,  the  anterior  one  rudimentary, 
the  four  developed  lobes  ending  in  acute  chitinized  points  and  each  bearing  a  subapical 
seta;  anterior  pair  of  lobes  more  widely  separated  and  bearing  on  lower  side  near  base  a 
small,  slender  lobule. 

Nepionotype. —  Ithaca,  New  York,  March  27,  1914. 

Neanotype. —  Ithaca,  with  type  larva,  April  9,  1914.     No.  2-1914. 

Paratypes. —  With  types. 

Ormosia  innocens  (O.  S.) 

1869     Rhyphoiophus  innocens  0.  S.      Mon.  Dipt.  N.  Amer.,  part  4,  p.  142. 

Ormosia  innocens  is   a   characteristic  early  spring  species,   the  adult 

flies  being  found  in  late  April  and  during  May.     On  May  12,  1917,  a 

few  pupae  in  an  advanced  stage  of  development  were  sifted  from  organic 

mud  from  Bool's  hillside,  Ithaca,  New  York.     One  of  these  emerged  as 

an  adult  on   the  following  day.     The  associated   crane-fly  larvae  that 

occurred  with  this  species  on  the  date  named  were  as  follows:  Bittaco- 

morphella  jonesi,    Dicranomyia  stulta,   Limnophila  adusta,  L.  fuscovaria, 

Ulomorpha    pilosella,     Penthoptera    albitarsis,     Rhaphidolabina    flaveola, 


926  CHARLES  PAUL  ALEXANDER 

Molophilus  hirtipennis,  Erioptera  megophthalma,  Ormosia  nigripila,  Tipula 
oropezoides,  T.  collaris,  T.  cayuga. 

Pupa. —  Length,  7.5-10.5  mm. 
Width,  d.-s.,  0.8-1  mm. 
Depth,  d.-v.,  1-1.2  mm. 

(The  smaller  measurements  are  those  of  males,  the  larger  those  of  females.) 

Head  and  thorax  brown;  mesonotum  before  declivity  darker  brown;  abdomen  pale  yellowish 
white,  the  sternal  and  tergal  sclerites  with  broad,  rectangular,  darker  areas,  producing  a 
banded  appearance;  breathing  horns  pale  yellow.  (In  older  pupae,  the  coloration  of  the 
head  and  the  thorax  is  much  darker.) 

Cephalic  crest  of  moderate  size,  bilobed;  viewed  from  side,  lobes  blunt,  with  two  lateral 
setae;  viewed  from  front,  lobes  separated  by  a  very  broad,  V-shaped  notch.  Between  antennal 
bases,  forehead  longitudinally  grooved,  these  lines  converging  between  eyes.  Labrum 
triangular,  subacute,  separating  the  small  labial  lobes.  Maxillary  palpi  of  moderate  length, 
tapering  to  blunt  apices.  Antenna  of  moderate  length,  reaching  to  just  beyond  wing  base. 
Pronotal  breathing  horns  rather  short  and  stout,  almost  straight  and  only  slightly  diverg- 
ing, subcylindrical,  compressed,  and  slightly  constricted  just  beyond  base.  Meso- 
notum precipitous,  carinate  medially;  at  crest  (Plate  LXX,  376)  with  numerous  pale 
hairs,  these  narrowly  interrupted  on  median  line;  lateral  angle  of  thorax  with  two  tiny  setae; 
a  strong  seta  slightly  dorsad  and  cephalad  of  wing  root;  about  four  small  setae  in  alinement 
across  mesonotum.  Wing  sheaths  moderately  broad,  ending  at  or  just  before  tip  of  second 
abdominal  segment.  Leg  sheaths  short,  tips  of  all  the  tarsi  ending  about  on  a  level,  or 
those  of  fore  legs  a  little  longer,  terminating  just  before  end  of  third  abdominal  segment. 

Abdominal  segments  divided  into  two  distinct  annuli  by  a  constriction  at  about  mid- 
length  of  segment;  sternum  and  tergum  of  posterior  ring  of  segments  2  to  8  near  caudal 
margin  with  large,  rectangular,  dusky  areas  which  appear  subchitinized;  caudal  margin  of 
this  area  with  a  fringe  of  long,  black  hairs,  there  being  about  seventy-five  of  these  on  inter- 
mediate segments;  on  pleural  membrane  a  very  small,  similar  area  bearing  from  twenty-five 
to  thirty-nine  hairs,  and  near  its  dorsal  margin  a  distinct  black  tubular  spiracle  on  segments 
2  to  7;  the  following  additional  setae  on  segments:  on  dorsal  segments,  rectangular  darkened 
areas  with  sparse  scattered  elongate  hairs  over  surface,  and  two  setae  on  each  anterior  ventral 
angle,  their  arrangement  oblique;  sternal  segments  similar,  but  the  two  setae  on  rectangular 
area  arranged  transversely;  just  caudad  of  ends  of  fringe  of  hairs,  one  or  two  isolated  hairs; 
pleural  membrane  opposite  basal  annulus  with  a  single  seta;  another  seta  opposite  posterior 
annulus  near  ventral  margin;  a  third  seta  on  ventro-cephalic  angle  of  pleural  setiferous 
area  just  above  level  of  spiracle;  segment  8  on  dorsum  with  five  pale,  gill-like  lobes  arranged 
in  a  quadrangle  or  a  pentagon,  the  anterior  lateral  pair  the  longest,  the  median  one  rather  the 
smallest;  at  base  of  anterior  pair,  two  setae  on  cephalic  face;  at  base  of  posterior  pair,  a 
single  seta  on  lateral  face.  Male  cauda  (Plate  LXX,  377  and  378)  terminating  in  two  very 
blunt  ventral  lobes  and  two  separated,  slender,  dorsal  lobes  projecting  caudad  and  dorsad. 
Female  cauda  similar  to  male  cauda,  but  tergal  acidothecae  very  elongate,  much  longer  than 
sternal  valves. 

Neanotype.—  Ithaca,  New  York,  May  12,  1917. 
Paratypes. —  Seven  pupae,  with  type. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  927 

Ormosia  nigripila  (O.  S.) 

1869     Rhypholophus  nigripilus  O.  S.     Mon.  Dipt.  N.  Araer.,  part  4,  p.  142. 

The  larvae  of  Ormosia  nigripila  are  common  in  rich  organic  mud  in 
shady  places.  Larvae  found  in  Needham's  Glen,  Ithaca,  New  York, 
on  March  27,  1914,  emerged  on  April  18.  Other  larvae  from  Coy  Glen, 
found  on  April  17,  emerged  on  May  1.  At  Orono,  Maine,  large  larvae 
were  found  on  July  14,  1913,  and  were  placed  in  rearing,  emerging  on 
the  26th  as  adult  males.  This  limits  the  pupal  duration  to  not  more  than 
two  weeks,  but  it  is  undoubtedly  much  less,  probably  not  more  than  a 
few  days  or  a  week. 

Pupa. —  Length,  5.2  mm. 

Width,  d.-8.,  0.9  mm. 
Depth,  d.-v.,  1  mm. 

Head,  thorax,  and  appendages  light  brown,  becoming  darker  in  maturity;  breathing  horns 
and  abdomen  pale  whitish. 

Cephalic  crest  with  lobes  small,  low,  and  rounded,  with  a  powerful  seta  at  tip,  directed 
forward.  Labrum  blunt  at  apex.  Labial  lobes  large,  subtriangular,  lateral  angles  obtusely 
pointed.  Sheaths  of  maxillary  palpi  stout,  rather  elongate.  Pronotal  breathing  horns 
(Plate  LXXII,  384)  rather  short  and  flat,  compressed,  slightly  expanded  beyond  base;  outer, 
or  posterior,  margin  with  about  five  small  tubercles,  at  least  one  of  which  is  setiferous. 
Mesonotum  very  steep  and  precipitous,  as  in  this  group  of  species;  crest  (Plate  LXXII,  383) 
tumid,  extensive,  on  either  side  with  abundant  coarse  yellow  bristles;  caudad  of  these,  four 
stout  setae  in  a  quadrangle,  two  on  either  side  of  median  line;  lateral  angles  of  thorax  with 
two  stout  setae;  a  strong  seta  just  above  wing  base.  Wing  sheaths  reaching  base  of  third 
abdominal  segment.  Leg  sheaths  modsrately  long,  extending  almost  to  midlength  of  fourth 
abdominal  segment;  tarsi  of  middle  legs  much  shorter  than  the  others,  hind  legs  a  very  little 
longer  than  fore  legs. 

Abdomen  with  segments  (Plate  LXXII,  385)  before  their  caudal  margin  bearing  transverse 
rows  of  slender  spines;  pleura  with  a  small  area  lying  a  little  cephalad  of  tergal  and  sternal 
rows  and  margined  behind  with  eight  to  ten  spines.  Spiracles  distinct,  tubular.  Setae 
as  follows:  on  pleura,  a  seta  on  anterior  annulus,  a  stout  seta  just  ventrad  of  spiracles, and 
a  third  lying  a  little  ventrad  and  cephalad  of  spiracle;  on  sternites,  two  stout  setae  near 
base  of  posterior  annulus;  on  tergites,  two  setae  on  posterior  annulus,  lying  transversely 
at  level  of  spiracle,  and  a  third  seta  just  cephalad  of  end  of  row  of  spines.  Male  cauda  (Plate 
LXXII,  386)  with  ventral  lobes  rather  slender,  narrowed  outwardly,  and  somewhat  pointed 
at  apex;  dorsal  lobes  curved  strongly  backward,  terminating  in  acute,  chitinized  points 
with  a  strong  seta  on  outer  face  before  tip  and  a  second  seta  nearer  base;  dorsum  of  segment  8 
with  five  brown,  chitinized  lobes  which  are  crowned  at  their  apices  with  a  circlet  of  spines; 
posterior  pair  the  longest  and  stoutest,  broad  at  base,  more  slender  outwardly,  with  a  large 
lateral  spine  and  about  three  smaller  inner  sp'nes;  anterior  pair  more  slender;  median  lobe 
slender,  crowned  with  a  circlet  of  about  six  small,  subequal  spines.  Female  cauda  (Plate 


928  CHARLES  PAUL  ALEXANDER 

LXXII,  387)  with  dorsal  terebra  elongate,  almost  straight  or  very  slightly  upturned;  eighth 
segment  with  dorsal  pentagon  of  spine-tipped  lobes  quite  as  in  male. 

Ormosia  meigenii  (O.  S.) 

1859     Erioptera  meigenii  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  226. 

Ormosia  meigenii  is  one  of  the  commonest  species  of  the  genus,  occurring 
in  small  dancing  swarms  in  early  spring.  Larvae  of  this  species  were 
taken  on  April  10,  1914,  in  organic  mud  from  Needham's  Glen,  Ithaca, 
New  York.  An  adult  female  emerged  on  May  4. 

Larva. —  Length,  6.5  mm. 

Diameter,  0.5-0.6  mm. 

Color,  light  yellow. 

Form  elongate,  terete.  Spiracular  disk  (Plate  LXX,  379)  about  as  in  0.  nubila,  dorso- 
median  lobe  the  smallest,  ventral  lobes  with  brown  lines  rather  close  together,  the  distal 
line  the  broadest;  lateral  lobes  with  lines  rather  short,  not  contiguous  at  their  distal  ends 
and  not  extending  past  midlength  of  spiracles;  dorso-median  lobe  with  marks  oval,  solidly 
dark  brown.  Lobes  fringed  with  moderately  long,  yellowish  setae,  which  are  almost  lacking 
at  extreme  tips;  disk  between  spiracles  unmarked.  Spiracles  large,  transversely  oval, 
ssparated  by  a  distance  a  little  greater  than  the  long  diameter  of  one. 

Head  capsule  and  mouth  parts  almost  as  in  0.  nubila;  mandible  with  the  long,  slender 
teeth  of  that  species. 

Pupa. —  (Described  from  a  cast  skin.) 

Cephalic  crest  consisting  of  low,  rounded  lobes,  each  with  a  seta  on  anterior  lateral  face. 
Pronotal  breathing  horns  of  moderate  length,  flattened,  a  little  narrowed  toward  tip,  outer 
margin  with  fine  tubercles.  Mesonotum  not  so  declivitous  as  usual  in  the  genus,  with  tiny 
roughenings  at  crest;  the  usual  two  setae  at  lateral  angle  of  thorax  present,  another  above 
each  wing,  and  four  more  in  a  transverse  row  at  level  of  axilla  of  wing;  a  seta  on  mesonotum, 
close  to  median  line,  just  below  crest.  Middle  legs  much  shorter  than  fore  and  hind  legs. 

Subterminal  rows  of  setae  on  abdominal  segments  consisting  of  slender,  acute  spines, 
with  a  few  setae  interspersed;  on  pleura  the  spines  few  in  number.  Spiracles  elongate, 
tubular.  Female  cauda  with  tergal  sheaths  very  long,  sternal  sheaths  short,  their  tips 
blunt;  on  dorsum  of  eighth  abdominal  segment  five  small  unarmed  tubercles,  which  are 
blunt  or  nearly  so,  anterior  pair  more  widely  separated  than  posterior  pair.  (In  the  shape 
of  the  cephalic  crest,  the  breathing  horns,  and  the  declivity  of  the  mesonotum,  this  species 
resembles  0.  nigripila;  but  the  arrangement  of  setae  on  the  abdomen,  and  the  great  reduction 
and  unarmed  condition  of  the  lobes  on  the  eighth  abdominal  tergite,  are  distinctive.) 

Nepionotype  —  Ithaca,  New  York,  April  10,  1914. 
Neanotype. —  Cast  pupal  skin,  Ithaca,  May  4,  1914. 

Genus  Helobia  St.  Farg.  et  Serv.  (Gr.  marsh  + 1  live) 

1825  Helobia  St.  Farg.  et  Serv.     Encyclop.  Method.  Ins.,  vol.  10,  p.  585. 

1830  Symplecta  Meig.     Syst.  Beschr.,  vol.  6,  p.  282. 

1865  Idioneura  Phil.     Verh.  Zool.-Bot.  Ges.  Wien,  vol.  15,  p.  615. 

1886  Symplectomorpha  Mik.     Wien.  Ent.  Zeitung,  vol.  5,  p.  318. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  929 

Larva. —  Form  moderately  elongated,  body  terete.  Spiracular  disk  surrounded  by  five 
subequal  lobes  which  are  marked  with  V-shaped  brown  lines,  in  some  specimens  the  inner 
faces  of  the  three  most  dorsal  lobes  being  entirely  brownish  black.  Anal  gills  moderately 
elongate.  Head  capsule  as  in  the  Eriopterini.  Antenna  with  the  apical  papilla  very  short, 
subpyriform. 

Pupa. —  Cephalic  crest  setiferous.  Pronotal  breathing  horns  elongate-cylindrical,  directed 
ventrad  and  cephalad,  with  rows  of  breathing  pores  along  dorsal  face.  Mesonotum 
declivitous,  at  broad  crest  armed  with  numerous  chitinized  spines;  an  arcuated  longitudinal 
row  of  six  small  pits  extending  from  wing  axil  toward  crest  of  thorax.  Wing  sheaths  ending 
before  tip  of  second  abdominal  segment.  Leg  sheaths  ending  about  opposite  tip  of  third 
abdominal  segment;  tarsi  of  hind  legs  the  longest,  those  of  middle  legs  the  shortest,  fore 
legs  intermediate  in  length.  Abdominal  segments  with  a  strong  sub  terminal  armature  on 
ventral  segments,  much  weaker  to  lacking  on  dorsal  segments;  lateral  spiracles  distinct  on 
segments  2  to  7;  dorsum  of  segment  8  with  five  blunt  lobes. 

Helobia  is  a  small  genus  (five  species)  of  common  and  sometimes  very 
widely  distributed  crane-flies.  The  only  Noith  American  species,  Helobia 
hybrida,  is  apparently  the  most  widespread  tipulid  known,  ranging  over 
practically  the  entire  Holarctic  region,  southward  in  the  mountains  to 
India,  and,  in  the  New  World,  to  Central  America.  The  immature 
stages  of  the  known  species  are  spent  in  moist  earth  near  water.  Bruch 
(in  litt.)  mentions  the  rearing  of  H.  macroptera  (Phil.)  in  Argentina. 

Helobia  hybrida  (Meig.) 

1804  Limonia  hybrida  Meig.     Klass.,  vol.  1,  p.  57. 

1818  Limnobia  punctipennis  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  1,  p.  147. 

1830  Symplecta  punctipennis  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  6,  p.  283. 

1848  Limnobia  cana  Walk.     List  Dipt.  Brit.  Mus.,  vol.  1,  p.  48. 

Helobia  hybrida  is  undoubtedly  the  most  widely  distributed  North 
American  crane-fly.  Beling  (1879:50-51)  found  larvae  and  pupae  at 
the  end  of  July,  1876,  in  wet,  sandy  earth  along  the  margins  of  small 
brooks  in  deciduous  woods.  Adults  emerged  in  his  breeding  cages  on 
July  27  and  August  6.  Hart  (1898  [1895] :  199-200)  found  the  larvae  in 
similar  sandy  situations  along  the  Illinois  River,  associated  with  the 
larvae  of  Tabanus  atratus  Fabr.  He  suggests  that  it  may  serve  as  food 
for  this  horse-fly  larva.  Larvae  of  Helobia  were  especially  abundant  on 
May  17,  and  these  transformed  to  adults  within  a  month.  Females 
were  observed  ovipositing  along  the  shore,  patting  the  valves  of  the 
ovipositor  against  the  moist  sand.  Malloch  (1915-17b:  229-230)  has 
given  additional  notes  on  the  structural  details  of  Hart's  material. 


930  CHARLES  PAUL  ALEXANDER 

Dr.  Adam  Boving  found  this  species  in  Iceland  and  made  careful  notes 
on  the  burrows  made  by  the  larvae.  Thru  the  kindness  of  Dr.  Boving, 
the  writer  is  able  to  include  a  translation  of  his  manuscript.  The  writer 
is  indebted  also  to  Dr.  Lundbeck,  director  of  the  museum  at  Copenhagen, 
for  the  loan  of  this  material  for  study.  These  are  the  specimens  discussed 
later  in  this  paper.  Boving' s  notes  were  made  at  Fell  Station,  southeast 
Iceland,  in  1908.  The  translation  follows: 

Inside  the  moraine  of  1877,  in  the  low  hnd  where  ice  was  standing  in  1886,  quantities  of 
dipterous  larvae  were  found  in  the  moist  sand  on  the  bottom  of  flat  hollows  which  at  times 
are  flooded  by  water  and  at  times  are  partly  drained,  as  was  the  case  on  the  day  when  the 
following  observations  were  taken. 

The  whole  dark,  moist  surface  of  the  bottom  was  covered  by  an  irregular  system  of  slightly 
elevated,  long,  tubular  galleries,  some  of  which  were  rather  straight,  some  formed  broken 
lines,  some  peculiar  arabesques,  and  some  plain  spirals.  The  width  of  the  galleries  was  about 
the  size  of  an  ordinary  pinhead,  some  a  trifle  larger,  some  a  little  smaller.  In  the  anterior 
part  of  each  gallery  was  found  either  a  cylindrical  white  tipulid  larva  (Helobia)  about  one 
centimeter  long,  or  another  dipterous  larva  of  the  same  general  size  and  appearance.  The 
larvae  were  found  just  below  the  surface.  It  was  not  always  easy  to  capture  them,  for  when 
I  pushed  my  knife  under  -the  mouth  of  the  gallery  they  moved  quickly  backward,  and  then, 
digging  deeper  into  the  soil,  made  a  new  gallery  that  branched  off  from  the  main  one.  It 
was  not  possible  to  distinguish  the  galleries  of  the  crane-fly  larvae  from  those  of  the  other 
dipterous  associate.  Very  often,  from  the  mouth  of  the  spiral  galleries,  one-third  of  a 
broken  pupal  skin  stuck  out;  but  larvae  were  found  also  in  many  of  these  galleries. 

The  imagines  of  the  two  Diptera  were  present  in  large  numbers,  some  flying  close  to  the 
ground,  others  resting  on  it.  Both  forms  were  long-legged  and  capable  of  running  over 
the  water  film.  I  secured  a  pair  of  both  in  copulation.  The  eggs  were  found  on  the  moist 
surface,  singly  or  in  small  masses  of  two  or  three  together. 

'  The  larvae  feed,  of  course,  on  organic  particles  in  the  sand.  The  imagines  were  not  observed 
to  take  any  nourishment  at  all;  they  copulated  as  soon  as  they  had  left  the  pupal  skins,  and 
I  did  not  find  them  in  any  other  place  than  on  the  bare,  moist  soil  where  the  larvae  lived; 
not,  for  instance,  on  flowers  growing  near  by. 

A  small  carabid  (probably  Bembidion  grapii  Gyll.)  was  present  in  the  locality  in  com- 
paratively large  numbers,  evidently  preying  on  the  larvae  of  the  Diptera.  A  single  cara- 
bid larva  also  was  found;  from  its  size  and  habitus  it  may  very  well  be  the  larva  of  the 
Bembidion. 

A  small  black  spider  was  probably  feeding  on  the  imagines  of  the  Diptera.  It  did  not  make 
a  regular  web,  but  spun  a  number  of  single  threads,  each  about  two  feet  long,  attaching  them 
to  a  piece  of  gravel  and  proceeding  from  this  as  a  common  center,  spreading  the  threads 
close  to  the  ground  like  radii,  and  finally  fastening  the  ends  to  small  grams  of  sand. 

The  adult  flies  of  Helobia  hybrida  are  very  common.  They  are  the 
first  tipulids  to  appear  on  the  wing  in  spring,  some  appearing  in  early 
March  or,  in  open  winters,  even  in  late  February.  They  remain  until 
late  in  the  fall.  The  writer  has  noted  the  females  running  about  on  the 
wet  sand  along  the  banks  of  the  Kaw  River  at  Lawrence,  Kansas,  and 
ovipositing  quite  as  described  by  Hart  (1898  [1895]:  199-200). 

Larva. —  Length,  7.8-10  mm. 

Diameter,  0.6-0.7  mm. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  931 

Coloration,  pale  brownish  yellow. 

Form  moderately  elongated.  Body  covered  with  a  sparse  appressed  pubescence.  Spi- 
racular  disk  (Plate  LXXIII,  388)  surrounded  by  five  lobes,  the  dorsal  one  notably  smaller 
than  the  paired  lobes;  inner  face  of  lobes  with  two  subparallel  brown  lines,  connected  at 
their  outer  ends  to  form  narrow  V's;  disk  marked  with  about  six  spots  between  spiracles, 
the  largest  at  base  of  ventral  lobes;  disk  fringed  with  short  hairs,  which  are  interrupted  for 
a  short  distance  between  lobes.  Anal  gills  moderately  elongated,  pale.  (Beling  describes 
the  three  dorsal  lobes  of  the  disk  as  having  the  entire  inner  face  shiny  blackish  brown,  and 
the  ventral  lobes  merely  margined  with  brownish;  there  would  thus  seem  to  be  some  varia- 
tion in  the  character  and  degree  of  markings  in  this  species.) 

Head  capsule  as  in  the  tribe.  Antenna  with  the  basal  segment  stout,  cylindrical,  the 
apical  papilla  very  short,  subpyriform.  Mandible  flattened;  teeth  large  but  very  bluntly 
rounded;  apical  tooth  the  largest,  with  a  slightly  smaller  denticle  en  either  side. 

Pupa. —  Length,  7-9  mm. 

Cephalic  crest  small;  lobes  pointed,  directed  ventrad,  each  bearing  a  short,  stout  seta. 
Labrum  narrow.  Labial  lobes  large,  divergent,  caudal  margin  almost  straight  across.  Sheaths 
of  maxillary  palpi  slender,  narrowed  at  tip  (Plate  LXXIII,  390).  Antenna  moderately 
angulated,  ending  just  beyond  base  of  wing.  Pronotal  breathing  horns  elongate-cylindrical, 
directed  ventrad  and  cephalad,  with  rows  of  breathing  pores  along  dorsal  face;  ventral  face 
transversely  wrinkled.  MesDnotum  moderately  declivitous,  at  broad  crest  (Plate  LXXIII, 
389)  armed  with  numerous  black  chitinized  spines  directed  backward;  these  spines  most 
num  rous  proximally,  interrupted  by  a  medianispace,  less  numerous  along  shoulder;  four  setae 
in  a  transverse  row  across  mesonotum,  two  on  either  side  of  median  line;  lateral  angle  of 
thorax  broad  and  blunt,  with  two  small  setae,  the  outermost  one  the  larger,  the  inner  one 
about  half  its  size;  an  arcuated  longitudinal  row  of  about  six  pits  extending  from  above 
axil  of  wing  toward  crest  of  mesonotum.  Wing  sheaths  short,  ending  before  tip  of  second 
abdominal  segment.  Leg  sheaths  ending  opposite  or  slightly  beyond  tip  of  third  abdominal 
segment;  hind  legs  much  the  longest,  middle  legs  much  the  shortest,  fore  legs  intermediate 
(Plate  LXXIII,  391). 

Abdomen  with  a  rather  strong  armature  of  stout  black  spines  on  sternal  segments,  on 
segment  3  this  appearing  as  a  small  area  of  about  ten  spines,  on  either  side  of  tips  of  hind 
tarsi;  tergal  armature  much  weaker  or  lacking;  lateral  spiracles  very  distinct,  tubular,  in 
cast  pupal  skin  the  principal  tracheal  trunks  being  very  conspicuous.  Female  cauda  (Plate 
LXXIII,  393)  with  tergal  valves  the  longest,  terminating  in  sharp  cylindrical  points;  sternal 
valves  much  shorter,  blunt  at  tips;  dorsum  of  segment  8  with  five  blunt  lobes.  Male  cauda 
(Plate  LXXIII,  392  and  394)  with  dorsal  valves  the  longest,  each  terminating  in  a  long, 
subacute,  chitinized  spine  directed  dorsad  and  laterad  and  bearing  before  its  tip  two  stout 
setae;  ventral  lobes  stout  and  blunt,  longer  than  dorsal  lobes. 

Nepionotype. —  Fell,  between  Oraefa  and  Heineberg,  southeastern  Iceland. 

Neanotype. —  Cast  pupal  skin,  with  type. 

Paratypes.—  Numerous  cast  pupal  skins,  with  types  (in  the  collection  of  the  Copenhagen 
Museum). 


932  CHARLES  PAUL  ALEXANDER 

Genus  Trimicra  Osten  Sacken  (Gr.  three  +  small) 

1861     Trimicra  O.  S.    Proc.  Acad.  Nat.  Sci.  Phila.,  p.  290. 

Trimicra  is  a  small  genus  of  crane-flies,  including  about  fifteen  described 
species  found  in  most  parts  of  the  world,  almost  all  being  forms  of  moderate 
size  and  obscure  coloration.  These  various  species  bear  a  close  resem- 
blance to  one  another  and  are  hard  to  distinguish  specifically. 

Bergroth  and  other  European  writers  consider  the  genus  Trimicra  as 
being  the  same  as  Psiloconopa  Zetterstedt,  but  at  this  time  the  writer 
is  not  entirely  willing  to  accept  this  view. 

Beling  (1879:48)  described  what  he  took  to  be  the  larva  of  one  of 
these  flies,  and  in  his  key  to  the  larvae  of  crane-flies  (1886:206)  he 
included  it  in  close  proximity  to  the  Pediciini.  As  mentioned  elsewhere 
in  this  paper,  it  is  highly  probable  that  Beling  described  a  pediciine  larva, 
but  by  an  accident  had  larvae  of  Trimicra  in  his  breeding  jars,  the  latter 
larvae  emerging  first  and  confusing  the  author. 

Gerbig  (1913:161-163)  describes  the  real  larva  of  Trimicra  pilipes 
(Fabr.),  the  best-known  species  of  the  genus.  It  is  found  along  the 
margins  of  flowing  streams  with  muddy  banks.  It  is  a  dark-colored 
larva,  about  15  millimeters  in  length  and  about  2  millimeters  in 
diameter.  The  skin  of  the  body  is  similar  to  that  of  the  larvae  in  the 
typical  subgenus  of  the  genus  Limnophila,  being  covered  with  chitinous, 
hairlike  projections,  which  in  Trimicra  are  longest  on  the  dorsal  surface  of 
the  body.  On  each  segment  there  are  solitary  elongate  bristles,  above  and 
below  each  of  which  is  a  gland.  The  spiracular  disk  (Plate  LXXIII,  395) 
is  surrounded  by  five  nearly  equal  lobes,  which  have  the  inner  faces 
marked  with  equal  blackened,  chitinized  areas.  On  the  lateral  margins 
of  each  lobe,  but  occupying  only  the  distal  part  of  the  lobe  and  not  con- 
tinuous around  the  disk,  is  a  fringe  of  moderately  long  hairs.  At  the 
tip  of  each  ventral  lobe  are  two  bristles,  and  at  the  tip  of  each  lateral 
lobe  is  a  single  bristle,  these  being  inserted  outside  the  line  of  hairs  and 
surrounded  by  a  bright  circular  area.  The  ventral  sensory  bristles  found 
in  some  crane-fly  larvae  (as  Tipula  variipennis)  are  lacking.  At  the 
base  of  the  lateral  lobes  are  the  spiracles,  which  are  generally  similar  to 
those  in  the  subgenus  Limnophila.  Gerbig  discusses  in  detail  the  structure 
of  the  spiracles,  the  felt  chamber,  and  the  musculation  of  this  region 
of  the  body. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  933 

Bruch  (in  Hit.)  mentions  the  rearing  of  Trimicra  reziproca  (Walk.)  in 
Argentina  from  larvae  very  similar  to  that  described  above,  occurring  in 
the  same  type  of  habitat. 

Genus  Empedomorpha  Alexander  (Gr.  Empeda  +  shape) 

1916     Empedomorpha  Alex.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  507-508. 

Empedomorpha  is  a  monotypic  genus  of  flies,  evidently  related  to 
Trimicra.  It  is  very  curious  in  its  marked  sexual  dimorphism,  the  male 
having  an  extremely  large,  hairy  stigma,  which  encroaches  upon  the 
adjoining  veins  and  often  distorts  them.  The  fly  is  still  very  insufficiently 
known.  It  is  a  prairie-inhabiting  species,  occurring  from  South  Dakota 
to  Texas  and  New  Mexico.  Nothing  is  known  concerning  the  immature 
stages,  and  the  following  observations  on  the  habits  and  occurrence  of 
the  adult  flies  are  all  that  are  available. 

Empedomorpha  empedoides  (Alex.) 

1916     (?)  Trimicra  empedoides  Alex.     Can.  Ent.,  vol.  48,  p.  44^5. 

Empedomorpha  empedoides^  as  stated  under  the  generic  account  above, 
is  a  prairie-inhabiting  species.  Adult  flies  were  found  running  about 
on  the  sand  flats  of  the  Arkansas  River,  near  Cimarron,  Kansas,  from 
July  13  to  15,  1917,  by  H.  L.  Fackler  and  the  writer.  The  vegetation 
of  the  sand  bars  here  is  very  sparse  to  almost  lacking,  a  few  psammophytic 
grasses  being  the  main  element.  Associated  with  these  flies  on  the  sandy 
surface  were  a  characteristic  group  of  sand-loving  insects,  of  which  the 
following  were  the  most  constant :  larvae  and  adults  of  tiger  beetles, 
representing  at  least  three  species  of  Cicindela;  ground  beetles,  Carabidae, 
including  .'such  genera  as  Omophron,  Dyschirius,  Bembidion,  Tachys,  and 
a  few  others;  rove  beetles,  Staphylinidae,  such  as  Stenus;  Heteroceridae, 
Psammocharidae,  Asilidae,  Saldidae,  and  similar  groups.  In  the  cool  of 
early  evening,  numerous  small  spiders  that  lurk  in  hollows  and  in  deserted 
insect  burrows  during  the  day  emerge  from  hiding  and  become  active. 

The  crane-flies  were  observed  during  the  hours  of  bright  sunlight,  when 
the  temperature  registered  over  100°  F.  in  the  sun.  They  run  rapidly 
over  the  moist  sand,  their  course  being  very  shifting  and  zigzag,  quite 
like  that  of  tiger  beetles.  They  fly  readily  but  only  for  short  distances, 
and  prefer  to  alight  on  the  sand  rather  than  on  the  vegetation.  They 
walk  awkwardly  over  the  sand,  but  are  able  to  crawl  up  grass  blades  or 


934  CHARLES  PAUL  ALEXANDER 

similar  objects.  The  habits  of  the  adult  flies  are  strikingly  like  those 
of  Helobia  (page  930).  The  writer  is  sure  that  the  larvae  are  to  be  found 
in  the  sand  in  these  same  haunts. 

Genus  Qnophomyia  Osten  Sacken  (Gr.  darkness  +  fly) 

1859  Gnophomyia  O.  S.  Proc.  Acid.  Nat.  Sci.  Phila.,  p.  223. 
1867  Furina  Jaenn.  Abhandl.  Senkenb.  Ges.,  vol.  6,  p.  318. 
1911  Dasymallomyia  Brun.  Rec.  Indian  Mus.,  vol.  6,  p.  304. 

Larva. —  Body  slender,  tapering  toward  ends,  with  transverse  welts  on  intermediate 
abdominal  segments.  Spiracular  disk  surrounded  by  five  subequal  lobes.  Anal  gills  con- 
sisting of  four  blunt,  rounded  lobes,  constructed  for  propulsion  rather  than  for  a  respiratory 
function.  Head  capsule  moderately  elongated,  rather  compact  for  the  Eriopterini.  Antenna 
rather  small,  apical  papilla  elongate-oval.  Mandible  slender,  with  a  long  apical  point  and 
three  teeth  along  ventral  cutting  edge. 

Pupa. —  Cephalic  crest  a  low,  blunt  tubercle  on  either  side  of  median  line,  each  tipped 
with  a  long  seta.  Pronotal  breathing  horns  small,  narrowly  trumpet-shaped.  Mesonotum 
moderately  declivitous,  at  crest  practically  unarmed;  a  strong  seta  at  lateral  angle  of  thorax 
and  another  on  either  side  of  mesonotum  behind  crest.  Wing  sheaths  short,  ending  before 
tip  of  second  abdominal  segment.  Leg  sheaths  very  short,  reaching  just  beyond  wings, 
attaining  end  of  second  abdominal  segment;  all  the  tarsi  ending  about  on  a  level.  Abdomen 
provided  with  long  setae;  spiracles  on  abdominal  segments  2  to  7. 

Gnophomyia  is  a  small  genus  including  about  forty  described  species 
which  are  most  abundant  in  the  Tropics  of  the  New  World.  The  European 
Gnophomyia  tripudians  Bergr.  has  recently  been  reared  by  Gamkrelidze 
(1913,  .a  and  b)  and  by  Keilin  (Edwards,  1919b).  The  former  found 
larvae  in  large  numbers  in  the  viscous,  semi-decomposed  mass  of  tissue 
beneath  the  bark  of  a  fallen  Carolina  poplar,  associated  with  Miastor 
metraloas  Meinert.  These  larvae  were  found  near  Paris,  France,  in 
March,  1911.  Gamkrelidze  records  a  gregarine  parasite  in  the  intestine 
and  a  nematode  worm  in  the  body  cavity.  The  species  was  later  reared 
in  England  from  dead  oak  by  Keilin,  who  has  discussed  and  figured 
glands  in  the  larvae  (1913:3).  The  only  American  species  that  has  been 
reared  is  the  common  Gnophomyia  tristissima,  discussed  later. 

Gnophomyia  rufa  Hudson,  of  New  Zealand,  has  recently  been  discussed 
in  some  detail  by  Hudson  (1920:32-33).  It  is  occasionally  found  in  dense 
forests  in  the  vicinity  of  Wellington.  The  larva  lives  in  and  feeds  on  the 
semi-liquid  vegetable  detritus  which  accumulates  in  large  quantities  at 
the  bases  of  the  leaves  of  Astelia  Solandri,  a  common  and  very  conspicuous 
epiphytic  plant  in  most  of  the  primitive  native  forests.  The  length  of 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  935 

the  fully  grown  larva  is  slightly  over  30  millimeters.  It  is  subcylindrical, 
considerably  flattened.  Oval  warts  armed  with  minute  teeth  are  situated 
on  both  surfaces  of  abdominal  segments  2  to  7.  The  larva  is  very  dark 
slaty  gray  in  color,  darker  toward  the  extremities.  The  spiracular  disk 
as  shown  by  Hudson's  colored  figure  is  very  small.  Apparently  only 
one  larva  inhabits  the  space  between  the  two  sheathing  leaves  of  the 
Astelia,  and  only  those  leaves  which  are  full  of  a  thick,  brown,  coffee- 
like  liquid  are  frequented.  The  pupa  is  inclosed  in  a  rather  tough, 
extremely  elongate,  silken  tube  situated  between  the  sheathing  leaves. 
It  rests  in  an  upright  position  in  the  midst  of  the  semi-liquid  mass.  The 
pupa  measures  about  38  millimeters  in  length,  being  very  elongate  with 
the  head  and  the  thorax  unusually  small.  The  two  pronotal  breathing 
horns  are  shaped  somewhat  like  a  bivalve  shell.  Abdominal  segments 
3  to  6  at  the  base  on  the  dorsal  side  have  finely- toothed  warts;  the  ventral 
surface  has  plain  ridges.  It  is  probable  that  the  present  species  is  not  a 
true  Gnophomyia,  but  until  more  is  known  of  this  species  and  its  relatives 
it  should  be  referred  to  this  genus. 

The  Gnophomyia  pilipes  referred  to  by  Beling  (1879:42)  and  by  Gerbig 
(1913:161-163)  pertains  to  Trimicra  (page  932). 

G.  tristissimah&s  been  recorded  by  Malloch(  191 5-17 b:  230-231)  as  living 
in  wet  mud,  but  this  is  an  error.  The  writer  has  material  from  exactly 
the  same  source  as  Malloch's,  received  from  James  A.  Hyslop,  and  this 
shows  that  the  haunt  of  the  larvae  is  beneath  the  decaying  bark  of  trees  — 
an  unusual  habitat  for  one  of  the  Eriopterini,  which  for  the  most  part 
live  in  damp  sand  or  earth  near  water.  The  rearing  of  this  species  in 
New  York  (by  Young),  Massachusetts  (by  Johnson),  Maryland  (by 
Hyslop),  Virginia  (by  Shannon),  Kansas  (by  Alexander),  Illinois  (by 
Malloch  and  Alexander),  and  Texas  (by  Mitchell),  leaves  no  question 
that  the  immature  stages  are  to  be  found  beneath  the  decaying  bark  of 
the  larger  hardwood  trees,  the  tulip  tree,  Liriodendron  Tulipifera  Linn., 
being  often  preferred. 

Onophomyia  tristissima  0.  S. 

1859    Gnophomyia  tristissima  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  224. 
Gnophomyia  tristissima  is   an  interesting  black  fly  with   conspicuous 
yellow  halteres.     It  is  common  and  widely  distributed  thruout  the  eastern 


936  CHARLES  PAUL  ALEXANDER 

United  States  and  Canada.  As  stated  above,  the  species  has  been  reared 
on  at  least  seven  different  occasions  in  as  many  States,  the  most  com- 
plete account  being  that  by  Hyslop,  whose  specimens  and  manuscript 
notes  were  kindly  placed  at  the  writer's  disposal.  His  data  on  this  species 
are  as  follows: 

May  14,  1914.  Wolfsville,  Maryland.  Under  the  bark  of  a  rotten  stump  of  a  tulip 
poplar  (Liriodendron)  on  the  roadside  near  Warrenfeltz  schoolhouse,  field  on  the  left  going 
to  town.  I  found  a  great  number  of  amber-yellow  dipterous  larvae  (three  in  alcohol),  and 
also  three  pupae  slightly  shortened  and  with  the  thorax  and  legs  "ferruginous  and  the  abdo- 
men pale  amber.  They  were  in  a  very  moist  nidus  of  rotted  inner  bark;  placed  in  rearing  in 
a  tin  box. 

May  16.     One  adult  emerged  today  (pinned);  pupal  case  in  alcohol. 

May  18.  Three  adults  emerged  today  (pinned);  pupal  cases  in  alcohol.  Observed 
emergence  of  one  adult.  The  swaying  motion  observed  in  Tipula  infuscata  was  not  observed, 
but  the  adult  simply  glided  straight  out  of  the  pupal  case  by  a  wavelike  contraction  of  the 
abdomen.  The  whole  emergence  took  only  about  eight  seconds.  A  larva  pupated. 

May  23.  Adult  emerged.  Pupal  stage  five  days.  Placed  the  remainder  of  the  pupae 
(all  had  transformed  from  larvae  to  pupae)  in  alcohol.  The  pupae  are  quite  active  and  move 
under  a  shelter  if  exposed.  Just  before  emerging,  the  pupa  takes  on  a  black  color  on  the 
thorax  and  smoky  yellow  on  the  abdomen. 

Shannon's  material  was  reared  from  pupae  taken  under  the  bark  of  a 
dead  tulip  tree  at  Dead  Run,  Fairfax  County,  Virginia,  on  May  5, 
1913.  A  larva  that  is  undoubtedly  this  same  species  was  found  beneath 
the  same  tree  on  April  17,  1913.  Johnson  bred  this  species  from  larvae 
found  beneath  bark  at  Riverside,  Massachusetts,  on  April  24,  1905. 
The  Texas  specimens  were  found  by  Mitchell  beneath  the  bark  of  cotton- 
wood  (Populus)  at  Victoria,  on  June  30. 

At  Lawrence,  Kansas,  in  1919,  the  writer  found  a  few  larvae  under 
the  bark  of  a  box  elder,  associated  with  the  following  dipterous  larvae : 
Pterocalla  strigula  Loew,  Lonchaea  laticornis  Meig.,  Phaonia  harti  Mall. 
These  associated  species  were  kindly  determined  by  Mr.  Malloch.  In 
1920  the  flies  were  bred  from  under  the  bark  of  several  deciduous  trees  at 
Urbana,  Illinois,  by  Mr.  Malloch  and  the  writer. 

Well-preserved  specimens  of  the  larvae  are  not  available  to  the  writer, 
and  the  following  description  is  taken  direct  from  Malloch  (1915-17  b: 
230-231) : 

Larva. —  Length,  9-11  mm.  Slender,  slightly  tapering  toward  both  extremities,  more 
decidedly  towards  the  cephalic.  Body  yellowish  testaceous,  covered  with  dense  decumbent 
pile. 

Head  [Plate  LXXIV,  39o]  more  compact  than  that  of  Helobia,  the  lateral  rods  stouter; 
antennae  very  small;  maxillae  large,  produced  beyond  the  apex  of  the  narrow  labrum,  the  palpi 
stout;  labium  not  chitinized;  mandibles  slender,  with  a  long  sharp  apical  tooth  and  about  three 
poorly  defined  teeth  along  the  lower  lateral  margin.  Locomotor  organs  consisting  of  rather 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  937 

broad  fusiform  areas  on  anterior  portion  of  abdominal  segments  except  basal  and  apical; 
hairs  along  margins  of  segmental  incisions  more  distinct  than  elsewhere  because  of  their 
bein-;  slightly  curved  upward;  apical  segment  with  five  processes,  their  structure  and  mark- 
ings as  in  figure  [Plate  LXXIV,  397];  anal  ventral  blood-gills  in  the  form  of  four  short 
rounded  protuberances. 

The  pupa  is  described  from  four  cast  skins,  kindly  presented  by 
Mr.  Hyslop: 

Pupa.- —  Length,  8-10  mm. 

Cephalic  crest  a  low,  blunt  tubercle  on  either  side  of  median  line,  each  tipped  with  a  long, 
stout  seta.  Labrum  broad,  elongate,  obtuse  at  tip,  completely  separating  triangular  labial 
lobes.  Sheaths  of  maxillary  palpi  moderately  slender,  tapering  gradually  to  tip.  Antennal 
sheaths  moderately  elongated,  extending  to  about  opposite  wing  root;  basal  segments  angulate. 
Pronotal  breathing  horns  (Plate  LXXIV,  399  and  400)  small,  trumpet-shaped,  very  flattened, 
median  area  lacking  and  hence  the  margins  contiguous;  just  proximad  of  breathing  horns 
a  large,  roughly  triangular  lobe  on  either  side,  immediately  behind  which  are  two  small 
setae.  Mesonotum  moderately  declivitous,  at  crest  rather  tumid,  but  unarmed,  with  a 
few  parallel  grooves  on  either  side  of  median  line;  lateral  angle  of  thorax  very  sharp,  before 
tip  with  two  setae,  one  very  powerful,  the  more  dorsal  one  abortive;  a  strong  seta  on  either 
side  of  mesonotum  behind  crest.  Wing  sheaths  short,  ending  just  before  tip  of  second 
abdominal  segment.  Leg  sheaths  (Plate  LXXIV,  398)  very  short,  ending  just  opposite  tip 
of  second  abdominal  segment  and  thus  projecting  but  slightly  beyond  wing  tips;  hind  legs 
a  little  the  shortest,  but  no  striking  difference  in  length  of  various  sheaths. 

Abdominal  segments  subdivided  into  a  narrow  basal  ring  and  a  much  broader  posterior 
ring;  abdominal  segments  on  dorsum  with  four  transverse  rows  of  tiny  but  stout  setae,  two 
on  basal  ring  and  two  on  posterior  ring,  one  being  subbasal,  the  other  subterminal,  in  position; 
starnum  with  only  the  subterminal  row  of  setae  present,  but  this  well  marked,  the  other 
rows  merely  vestigial;  on  either  side  of  dorsum,  just  cephalad  of  ends  of  subterminal  row 
of  setae,  a  powerful  bristle;  in  alinement  with  these  and  subequally  spaced,  two  smaller 
setae;  a  strong  pleural  seta  on  a  raised  papilla  opposite  basal  ring  and  three  opposite  posterior 
ring,  the  two  anterior  being  larger,  the  posterior  one  very  small;  .sternum  with  a  strong  seta 
on  extreme  lateral  margin  of  posterior  ring;  lateral  spiracles  distinct,  on  segments  2  to  7. 
Male  cauda  (Plate  LXXIV,  401)  with  ventral  lobes  small,  blunt,  rather  widely  separated 
basally  but  converging  apically;  dorsal  lobes  powerful,  divergent,  and  rather  acute  at  tips; 
at  base  of  each  near  lateral  margin  a  short  bifid  knob  sending  one  arm  dorsad,  the  other 
laterad;  at  base  on  cephalic  angle  a  short,  stout  seta;  two  long,  powerful,  lateral  setae  on 
either  side,  and  a  single  powerful  seta  on  either  side  of  dorsum,  immediately  behind  which 
is  a  blunt  tubercle.  Female  cauda  with  sternal  valves  elongate,  powerful,  at  their  tips  ter- 
minating in  slender,  divergent  points;  dorsal  valves  small,  blunt,  divergent,  located  at  base 
of  sternal  valves,  at  their  tips  with  a  short,  slender  spine  directed  backward;  base  of  seg- 
ment about  as  in  male. 

Neanotype.—  Wolfsville,  Maryland,  May  16,  1913.     No.  234. 

Paratypes. —  Two  male  and  one  female  pupae  from  type  locality,  May  18,  1913. 


938  CHARLES  PAUL  ALEXANDER 

Genus  Qonomyia  Meigen  (Gr.  angle  +  fly) 

1818     Gonomyia  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  1,  p.  146. 
1850     Taphrosia  Rond.     Dipt.  Ital.  Prodr.,  vol.  1,  p.  182. 
1869    Goniomyia  O.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  176. 

Larva. —  Form  elongate,  terete.  Spiracular  disk  surrounded  by  five  blunt  lobes  which  are 
heavily  marked  with  brown,  in  some  species  (G.  alexanderi)  the  brown  suffusing  the  disk 
between  the  spiracles.  Head  capsule  of  eriopterine  type.  Mandible  with  lateral  teeth 
slender,  flattened.  Antenna  with  apical  papilla  elongate-oval.  Mentum  not  chitinized. 

Pupa.- —  Cephalic  crest  blunt,  the  surface  with  minute  roughenings.  Pronotal  breathing 
horns  flattened,  fanlike  (G.  sulphurella) ,  or  short,  trumpet-shaped.  Mesonotum  declivitous, 
at  crest  with  an  interrupted  transverse  row  of  six  to  eight  tubercles  which  are  densely  beset 
with  sharp  black  spicules.  Wing  sheaths  attaining  base  of  third  abdominal  segment.  Leg 
sheaths  moderately  elongated,  reaching  base  of  fourth  abdominal  segment;  tips  of  middle 
tarsi  ending  a  short  distance  before  apices  of  other  tarsi.  Armature  of  abdominal  segments 
weak.  Lateral  spiracles  distinct,  tubular.  Five  blunt,  fleshy  lobes  on  dorsum  of  eighth 
abdominal  segment. 

Gonomyia  is  a  largo  and  diverse  genus  of  small  crane-flies  (including 
more  than  one  hundred  known  species)  described  from  all  parts  of  the 
world.     They  are  divided  into  four  recent  subgenera,  of  which  three  — 
Gonomyia  Meig.,  Progonomyia  (new   name   for  Gonomyella   Alex.,  pre- 
occupied), and  Leiporieura  Skuse  —  occur  in  the  Nearctic  fauna. 

The  immature  stages  of  the  known  species  are  spent  in  moist  sand  or 
earth,  usually  near  water.  In  Europe,  G.  tenella  Meig.  (Beling,  1879:56, 
mention  only)  was  found  in  August  in  damp,  sandy  earth  along  the  margin 
of  a  dried-up  brook. 

The  writer  has  found  the  immature  stages  of  Gonomyia  (Leiponeura) 
alexanderi  and  G.  (G.)  kansensis  in  wet  sand  near  rivers.  G.  sulphurella 
and  G.  subcinerea  O.  S.  occur  in  muddier  and  more  stagnant  conditions 
near  ponds  and  small  streams. 

Not  enough  larvae  arc  available  for  study  to  require  a  key  at  this 
time.  The  pupae  of  the  known  Nearctic  species  may  be  distinguished  by 
the  following  key: 

1.  Pronotal  breathing  horns  narrow  at  base,  expanded  distdly  into  a  very  flattened,  fan- 

like  blade  with  delicate  and  anastomosing  nervures G.  sulphurella  O.  S.  (p.  €40) 

Pronotal  breathing  horns  not  as  above,  more  earlike  or  trumpet-shaped 2 

2.  Pronotal   breathing   horns   massive,    trumpet-shaped;    literal   margin   of    thorax   before 

wing  root  produced  into  an  angle;  male  cauda  small,  elongate,  dorsal  lobes  a  little 
shorter  than  ventral  lobes,  with  two  stout  lobes  on  dorsal  side  far  removed  from  their 

bass G.  alexanderi  (Johns.)  (p.  939) 

Pronotal  breathing  horns  flattened,  earlike  or  narrowly  trumpet-shaped;  lateral  margins 
of  thorax  above  wing  root  broad  and  blunt;  male  cauda  short,  stout,  dorsal  and  ventral 
lobes  subequal  in  length,  the  latter  closely  approximated  along  median  line,  the  former 
widely  separated,  at  their  base  with  two  acute  points G.  kansensis  Alex.  (p.  941) 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  939 

(Subgenus  Leiponeura  Skuse) 

1889    Leiponeura  Skuse.     Proc.  Linn.  Soc.  N.  S.  Wales,  ser.  2,  vol.  4,  p.  795. 
1915    Lipophleps  Bergr.     Psyche,  vol.  22,  p.  55. 

Gonomyia  (Leiponeura)  alexanderi  (Johns.) 

1912     Elliptera  alexanderi  Johns.     Psyche,  vol.  19,  p.  3. 

The  beautiful  crane-fly  Gonomyia'  alexanderi  is  locally  common  in  the 
eastern  United  States.  The  adult  flies  may  be  swept  from  rank  vegetation 
in  the  neighborhood  of  streams.  When  resting,  the  adults  have  a  char- 
acteristic position,  the  fore  legs  st'anding  straight  ahead  and  almost 
parallel,  the  middle  legs  extended  laterally  and  slightly  forward,  the 
hind  legs  directed  backward  but  widely  divergent,  and  the  wings  folded 
over  the  back.  This  is  the  characteristic  resting  position  for  the  genus. 
The  larvae  were  found  in  some  numbers  in  rather  coarse  sand,  around 
small  pools  of  water  near  the  Sacandaga  River,  Fulton  County,  New  York, 
on  June  5,  1914.  The  adults  emerged  on  June  16,  giving  a  pupal  period 
of  not  more  than  eleven  days  and  presumably  much  less.  The  description 
and  figures  of  the  pupa  are  made  from  the  cast  pupal  skin  of  the  male. 

Larva. —  Length,  8.3  mm. 

Diameter,  0.4-0.5  mm. 

Coloration  very  pale  yellow  or  yellowish  white. 

Form  terete,  elongated,  slender.  Body  with  a  sparse,  pale  pubescence,  at  posterior  margins 
of  segments  with  a  transverse  erect  ridge  of  stiff  hairs.  Spiracular  disk  (Plate  LXXV,  403) 
large,  flattened,  almost  pentagonal  in  outline,  surrounded  by  five  lobes;  dorso-medi^n 
lobe  small,  slender;  paired  lobes  very  short  and  blunt;  margin  between  lobes  almost  straight 
or  but  feebly  concave;  when  disk  is  partly  closed,  lobes  appearing  a  little  more  prominent; 
ventral  lobes  a  little  larger  than  lateral  lobes;  lobes  heavily  suffused  with  brown;  on  ventral 
lobes  a  lateral  dark  brown  line  running  dorsad  to  near  spiracles,  at  its  dorsal  end  connected 
across  disk  by  a  paler  brown  suffusion;  proximal  stripes  of  ventral  lobes  shorter  and  paler, 
above  their  inner  ends  with  a  small  brown  spot;  lateral  lobes  almost  entirely  suffused  with 
brown,  this  entirely  surrounding  spiracles  and  in  some  specimens  entirely  suffusing  disk 
between  spiracles,  this  mark  bifid  at  its  distal  end;  dorsal  lobe  indistinctly  marked  with 
very  pale  brown;  disk  margined  with  short,  pale  hairs  which  are  not  interrupted  and  are 
only  a  little  longer  at  tips  of  lobes.  Spiracles  widely  separated,  the  distance  between 
them  being  three  or  four  times  diameter  of  one;  spiracles  yellow,  centers  pale  brown. 

Head  capsule  as  in  the  tribe,  the  ventral  bars  broader  than  the  slender  dorsal  bars,  their 
inner  ends  not  expanded  or  toothed  to  form  the  mental  plate.  Labrum-epipharynx  moder- 
ately elongate,  densely  hairy.  Mentum  not  chitinized;  hypopharyngeal  region  a  cushion, 
covered  with  delicate,  short  setae.  Antenna  as  in  this  tribe,  basal  segment  moderately  elon- 
gate, densely  hairy,  apical  papilla  rather  small,  elongate-oval.  Mandible  (Plate  LXXV,  402) 


940  CHARLES  PAUL  ALEXANDER      » 

moderately  large;  apical  point  elongated,  slender;  ventral  cutting  edge  with  three  flattened 
teeth,  gradually  smaller  from  outermost  toward  base;  outermost  tooth  flattened,  a  little 
enlarged  distally,  about  as  long  as  apical  point;  basal  tooth  small,  acute;  a  prosthecal  append- 
age with  a  brush  of  hairs  beneath  it. 

Pupa.- —  Length  of  cast  pupal  skin,  about  5.5  mm. 

Cephalic  crest  of  two  prominent  lobes,  blunt  at  tips,  their  surface  granulated.  Labial 
lobes  blunt  at  tips.  Sheaths  of  maxillary  palpi  moderately  stout,  tapering  suddenly  to  sharp 
apices.  Antenna  with  basal  segment  very  angulated.  Pronotal  breathing  horns  (Plate 
LXXVI,  408  and  409)  massive,  short,  trumpet-shaped,  flattened  laterally,  and  here  margined 
with  an  elevated  ridge,  along  which  are  scattered  the  rows  of  breathing  pores;  mouth  of  this 
trumpet  wide.  On  thorax  between  breathing  horns,  large,  rounded  lobes  which  are  minutely 
granulated.  Declivity  of  mesonotum  (Plate  LXXVI,  407)  somewhat  precipitous,  at  the 
rather  narrow  crest  with  about  six  small  tubercles  which  are  densely  beset  with  spicules; 
along  shoulder  a  similar,  but  more  elongate,  transverse  welt;  lateral  margin  of  thorax  before 
wing  root  projecting  out  as  a  sharp  angle  with  a  seta  at  its  base.  Wing  sheaths  reaching 
end  of  second  abdominal  segment.  Leg  sheaths  moderately  long,  attaining  base  of  fourth 
abdominal  segment;  fore  legs  a  very  little  longer  than  hind  legs;  middle  legs  much  shorter, 
ending  just  beyond  base  of  last  tarsal  segment  of  fore  legs. 

Abdominal  segments  divided  into  two  narrow  basal  rings  and  a  much  broader  posterior 
annulus.  Armature  of  abdomen  very  weak.  Male  cauda  (Plate  LXXVI,  410  and  411)  small, 
elongate;  ventral  lobes  a  little  longer  than  the  short,  blunt  dorsal  lobes;  on  dorsal  face  near 
end  of  eighth  segment,  two  stout  lobes  pointed  at  the  tips  which  are  directed  dorsad  and 
slightly  caudad;  eighth  segment  with  a  close  pentagon  of  pale,  slender  lobes,  the  posterior 
pair  larger  and  closer  together  than  the  anterior  pair,  the  median  lobe  the  smallest. 

Nepionotype. —  Sacandaga  River,  Fulton  County,  New  York,  June  5,  1914. 
Neanotypc. —  With  type  larva. 
Paratypes. —  Two  larvae  with  type. 

(Subgenus  Gonomyia  Meigen) 

Gonomyia  (Gonomyia)  sulphurella  O.  S. 

1859     Gonomyia  sulphurella  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  230. 

1869     Goniomyia  sulphurella  0.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  180-181. 

Gonomyia  sulphurella  is  a  handsome  little  crane-fly  which  is  very 
common  and  widely  distributed  thruout  the  eastern  and  central  United 
States.  Larvae  are  not  infrequent  in  mud  along  the  banks  of  streams. 
The  writer  has  bred  this  species  from  larvae  sifted  from  sandy  mud  from 
the  banks  of  Cascadilla  Pond,  Ithaca,  New  York,  where  they  occur 
associated  with  numerous  larvae  and  pupae  of  a  tabanid  (Chrysops  indus 
O.  S.),  a  stratiomyiid  (Odontomyia  sp.),  and  other  forms.  Larvae  collected 
on  May  14,  1913,  emerged  as  adults  on  June  1.  Adults  have  been  reared 
as  late  as  October  19  by  E.  A.  Richmond. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  941 

Pupa. —  Length  of  cast  pupal  skin,  about  6.5-7  mm. 

Cephalic  crest  small,  blunt.  Labrum  triangular,  apex  obtusely  rounded.  Labial  lobes 
subcircular,  outer  margin  rounded.  Sheaths  of  maxillary  palpi  short  and  stout,  at  apex 
suddenly  narrowed  (Plate  LXXVII,  414).  Antennal  sheaths  angulated  at  segments,  the 
organ  extending  to  beyond  base  of  wing.  Pronotal  breathing  horns  (Plate  LXXVII,  413) 
with  extreme  base  expanded,  the  neck  short,  constricted,  soon  passing  into  a  greatly 
expanded  and  very  compress3d  disk,  the  whole  suggesting  a  fan  or  certain  polypores;  margin 
of  this  disk  entire  or  gently  crenulated,  and  sloping  from  ventral  side  outward;  surface  finely 
nerved  and  reticulated.  Mesonotum  somewhat  precipitous,  crest  (Plate  LXXVII,  412) 
tumid,  with  rounded  knobs  arranged  transversely  along  it,  there  being  about  eight  isolated 
knobs  and  a  more  elongate  one  along  shoulders,  these  knobs  covered  with  minute  blackened 
spicules.  Lateral  margin  of  thorax  above  wing  root  forming  almost  a  right  angle.  Wing 
sheaths  ending  opposite  base  of  second  abdominal  segment.  Leg  sheaths  comparatively 
short,  ending  opposite  base  of  fourth  abdominal  segment;  fore  legs  a  little  the  longest,  middle 
legs  conspicuously  shorter  than  the  others. 

Abdominal  segments  divided  into  two  narrow  basal  rings  and  a  broad  posterior  annulus. 
Armature  of  abdominal  segments  weak,  posterior  ring  with  a  narrow  row  of  small  black 
spines  before  caudal  margin;  on  basal  ring  a  broad  band  of  microscopic  scabrous  points 
arranged  in  interrupted  transverse  rows,  there  being  about  seven  or  eight  of  these  rows 
to  a  band.  Spiracles  small  but  distinct.  Female  cauda  with  tergal  valves  short,  but  little 
longer  than  sternal  valves,  slightly  upturned,  ending  in  short,  cylindrical  tips;  near  base 
with  a  small,  blunt  tubercle  on  either  side;  dorsum  of  segment  8  with  five  rather  long,  pale 
lobes,  the  anterior  pair  more  slender  and  more  widely  separated  than  the  posterior  pair,  which 
are  sometimes  closely  approximated. 

Neanotype.—  Cast  pupal  skin,  Ithaca,  New  York,  October  19,  1915. 
Paratypes. —  Three  pupae  with  type. 

Gonomyia  (Gonomyia)  kansensis  Alex. 

1918  Gonomyiz  kansensis  Alex.  Can.  Ent,  vol.  50,  p.  158-160. 
Gonomyia  kansensis  is  a  prairie  species  of  the  cognatella  group  and 
appears  to  be  common  along  the  Arkansas  and  Kaw  Rivers  in  Kansas. 
Living  pupae  were  found  at  Larned  on  August  1,  1917,  in  sand  along 
the  banks  of  the  Arkansas  River,  where  they  were  associated  with  the 
typical  sand-loving  fauna,  including  Gelastocoridae,  Saldidae,  and  Carab- 
idae  (Omophron,  Dyschirius,  Bembidion,  and  Tachys).  The  observation 
of  a  small  ant  preying  on  a  living  pupa  of  this  fly  is  discussed  on  page 
729. 

Pupa.- —  Length  of  cast  pupal  skin,  6  mm. 

Similar  to  G.  alexanderi  in  moet  essentials  but  showing  the  following  differences:  anten- 
nal  sheaths  strongly  angulate  at  segments;  a  tubercle  on  ventral  face  of  antenna  at  base,  and 
another  at  inner  cephalic  margin  of  eye. 


942  CHARLES  PAUL  ALEXANDER 

Pronotal  breathing  horns  (Plate  LXXVII,  415  and  416)  flattened,  earlike  or  very  narrowly 
trumpet-shaped,  with  a  thick  marginal  ridge,  the  disk  restricted.  Lateral  angle  of  thorax 
above  wing  root  very  broad  and  blunt,  setiferous.  Leg  sheaths  of  fore  and  hind  legs  almost 
on  a  level,  those  of  middle  legs  shorter,  ending  a  little  beyond  midlength  of  last  tarsal  segment 
of  fore  legs.  Male  cauda  (Plate  LXXVII,  417  and  418)  with  dorsal  and  ventral  lobes  very 
short  and  blunt,  subequal  in  length;  ventral  lobes  closely  approximated  on  median  line; 
dorsal  lobes  widely  separated  at  their  base,  with  two  small  acute  points  directed  strongly 
dorsad,  divergent  at  their  tips,  each  with  two  small  setae  on  outer  face  near  base;  dorsum  of 
segment  8  with  five  lobes,  the  anterior  pair  a  little  more  widely  separated  than  the  posterior 
pair,  the  median  lobe  slender. 

Neanotype. —  Lamed,  Kansas,  August  1,  1917. 

Genus  Rhabdomastix  Skuse  (Gr.  rod  4-  whip) 

1889    Rhabdomastix  Skuse.     Proc.  Linn.  Soc.  N.  S.  Wales,  ser.  2,  vol.  4,  p.  828-829. 

The  genus  Rhabdomastix  includes  nearly  a  dozen  species,  some  of 
which  have  been  previously  described  as  Gonomyiae. 

The  immature  stages  of  Rhabdomastix  schistacea  (Schum.)  were  found 
by  Beling  (1886:195)  in  wet  earth  beside  a  stream  in  beech  woods  on 
May  6.  The  larva  measures  6  millimeters  in  length;  the  greatest  diameter 
is  0.8  millimeter.  The  body  is  strongly  dilated  in  the  anterior  part  and 
gradually  narrowed  behind.  The  integument  is  deep  brownish  yellow. 
The  spiracular  disk  is  short  and  blunt,  and  has  four  very  small,  tuberculate 
teeth,  the  lateral  pair  lying  somewhat  more  cephalad  than  the  more 
powerful  ventral  pair;  spiracles  small,  circular,  yellowish  brown,  separated 
by  a  distance  about  equal  to  four  times  the  diameter  of  one.  The  pupa 
has  the  mesonotal  declivity  provided  with  an  interrupted  crossrow  of 
small,  unequal,  chitinized  teeth. 

(Subgenus  Sacandaga  Alexander) 

1911     Sacandaga  Alex.     Ent.  News,  vol.  22,  p.  349-352. 

Rhabdomastix  (Sacandaga)  flava  (Alex.) 

1911    Sacandaga  flava  Alex.    Ent.  News,  vol.  22,  p.  351-352. 

Rhabdomastix  flava 'is  a  curious  fly  which  is  apparently  related  to 
Gonomyia  but  represents  a  quite  different  offshoot  of  the  Eriopterini. 
The  writer  believes  that  the  larvae  might  be  found  in  moist  earth  along 
streams,  but  at  present  they  are  quite  unknown.  The  following  notes 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  943 

een  PUbHshed  (Alexander, 


IQl 

number  of  individuals  participating  in  the  swarm  was  about  twenty  %tW  species  swam 
mg  nearby  at  the  same  time  were  Chironomus  hyperboreus,  var.  £rid*mfa  Joh  anTthe 
mayflies,  Ephemerella  exerucians  Walsh,  and  Siphlonisca  aerodromia  Ndm. 


Genus  Trentepohlia  Bigot  (named  after  J.  J.  Trentepohl) 

1854     Trentepohlia  Biojot.     Ann.  Soc.  Ent.  France  p  474 

Mongomioides  Brun.     Rec.  Indian  Mus.,  vol  6  p  296 
912     Mongomella  Enderl.     Zool.  Jahrb.,  vol.  32,  part  1,'  p.  61. 

Trentepohlia  is  a  tropicopolitan  genus  including  about  fifty-five  described 
species  arranged  in  six  subgenera  —  Trentepohlia  Bigot,  Anchimongoma 
Brun.,  Mongoma  Westw.,  Plesiomongoma  Brun.,  Paramongoma  Brun. 
and  Neomongoma  Alex.     Of  these  species,  fifteen  are  American  and  the 
remainder  are  Old  World  forms.     They  are  almost  all  species  of  delicate, 
ethereal  structure,  with  long,  slender  legs.     It  is  an  interesting  fact  that 
Trentepohlia  (Mongoma)  pennipes  has  been  observed  by  Jacobson  (De 
Meijere,  1911:50,  and   Edwards,  1912-13:211)  to  form   chains  on  hori- 
zontal spider  webs,  as  is  noted  herein  for  Thrypticomyia  (page  712)  and 
somewhat   similarly  for  Oropeza  (page  982).     H.    K.   Munro  has  sup- 
plied (in  litt.}  the  following  interesting  notes  on  the  habits  of  Trentepohlia 
(Trentepohlia)  humeralis  Alex,  as  observed  in  eastern  Transvaal  at  the 
end  of  April,  1920: 

Very  inconspicuous  when  flying  and  resting.  When  flying  resembles  very  much  a  small 
piece  of  thistledown.  Invariably  settles  on  under  side  of  twigs,  leaves,  and  similar  objects. 
Usually  found  among  bushy  undergrowth,  but  also  in  grass.  When  at  rest  the  wings  are 
lolded  along  back;  very  often  on  settling  the  insect  moves  itself  up  and  down  in  the  manner 

the  long-legged  harvest  spiders  (Phalangiidae).     Slow  flier. 

I  A  fossil  Trentepohlia,  T.  cruciferella  (Ckll.),  has  been  described  from  the 
Gurnet  Bay  Oligocene  (Cockerell,  1917b:  373-374).  Observations  on  the 
immature  stages  of  three  species  are  available. 


944  CHARLES  PAUL  ALEXANDER 

(Subgenus  Mongoma  Westwood) 

Trentepohlia  (Mongoma)  pennipes  (O.  S.) 

1887    Mongoma  pennipes  O.  S.     Berl.  Ent.  Zeit,  vol.  31,  part  2,  p.  204. 

The  immature  stages  of  Trentepohlia  pennipes  have  been  described  by 
De  Meijere  (1911:50-51)  as  follows:  Jacobson  found  the  larvae  at 
Semerang,  Java,  in  January,  190G,  in  decaying  plant  stems.  The  only 
larva  sent  was  9  millimeters  long  and  almost  1  millimeter  in  diameter, 
of  cylindrical  form,  only  slightly  narrowed  behind  and  brownish  in  color. 
The  head  capsule  was  entirely  retracted.  The  entire  body  was  thickly 
beset  with  fine,  short,  appressed  hairs;  in  addition  to  these,  on'the  ventral 
side  of  -each  of  the  six  intermediate  segments  were  transverse  swellings 
where  the  hairs  were  shorter  and  even  more  numerous.  Surrounding 
the  anus  were  four  long,  cylindrical,  anal  gills,  each  constricted  in  three 
or  four  places;  if  bent  forward,  the  anterior  pair  would  reach  the  middle 
of  the  penultimate  segment  of  the  body,  the  posterior  pair  being  somewhat 
shorter.  The  last  segment  of  the  body  was  truncated,  the  lower  angles 
being  somewhat  produced  and  provided  with  a  few  somewhat  longer 
hairs;  the  spiracles,  situated  in  the  upper  part  of  the  spiracular  field, 
were  relatively  small  and  somewhat  elongated. 

The  pupa  (Plate  LXXVIII,  419)  was  about  9  millimeters  long,  elon- 
gate, of  a  yellowish  brown  color,  the  abdomen  for  the  most  part  brighter. 
The  thorax  was  almost  smooth,  with  only  a  few  short,  brownish  yellow 
bristles.  The  abdomen,  except  on  the  anterior  S3gments,  was  thickly  set 
with  numerous  tubercles.  The  apex  of  the  abdomen  had  two  short,  thorn- 
like  projections,  curved  outwardly;  beneath  these  were  two  shorter 
tubercles,  and  four  short  tubercles  formed  a  quadrangle  on  the  dorsum 
of  the  last  segment.  The  pronotal  breathing  horns  consisted  of  flat- 
tened, leaflike  lobes,  the  upper  surface  of  which  was  scaly. 

(Subgenus  Paramongoma  Brunetti) 

Trentepohlia  (Paramongoma)  bromeliadicola  (Alex.) 

1912    Mongoma  bromeliadicola  Alex.    Ent.  News,  vol.  23,  p.  415-417. 

Trentepohlia  bromeliadicola  and  T.  leucoxena  have  a  larval  habitat 
which  has  not  been  found  elsewhere  in  the  family.  They  live  in  the 
water  that  gathers  in  the  leaf  axils  of  tropical  bromeliaceous  plants, 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  945 

/!      ^   ! 

spending  their  immature  stages  in  this  habitat,  where  they  are  associated 
with  a  remarkable  fauna  of  other  organisms.  The  adult  females  hav.e 
the  valves  of  the  ovipositor  greatly  elongated,  and  the  writer  has  sug- 
gested elsewhere  that  this  may  be  an  adaptation  for  laying  the  eggs  in 
this  habitat.  T.  bromeliadicola  was  reared  in  Costa  Rica  by  Picado, 
whose  important  paper  (Picado,  1913)  on  the  bromeliaceous  epiphytes 
contains  colored  figures  of  the  larva,  the  pupa,  and  the  adult.  From 
this  paper  it  is  seen  that  the  larva  (page  356,  figure  A,  and  plate  13, 
figure  4,  of  reference  cited)  is  not  unlike  that  of  T.  pennipes,  described 
above,  the  four  prominent  anal  gills  of  T.  bromeliadicola  (Plate  LXXVIII, 
420,  of  this  paper)  being  a  notable  feature  in  common,  altho  here  the 
constrictions  are  very  numerous,  there  being  twenty-five  or  thirty  shown 
in  the  figure.  The  pupa  (Picado,  1913:357,  fig.  51,  and  pi.  13,  fig.  2) 
has  the  pronotal  breathing  horns  (Plate  LXXVIII,  421,  of  this  paper) 
approximated  on  the  median  line,  and  the  sheaths  of  the  ovipositor  (Plate 
LXXVIII,  423)  greatly  elongated  to  contain  the  elongated  terebra  of  the 
adult  within.  According  to  Keilin  (1913),  the  tegumentary  glands  of  this 
larva  are  a  provision  against  drought,  which  is  the  great  source  of  danger 
to  organisms  living  in  this  habitat. 

Trentepohlia  (Paramongoma)   leucoxena  (Alex.) 

1915    Mongoma  leucoxena  Alex.     Ent.  News,  vol.  26,  p.  29-30. 

Trentepohlia  leucoxena  was  reared  by  Knab  in  Mexico,  from  larvae 
found  living  in  bromeliaceous  plants  quite  as  in  the  preceding  species. 

Genus  Teucholabis  Osten  Sacken  (Gr.  weapons  +   forceps) 
1859     Teucholabis  0.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  222. 

Larva. —  Form  elongate,  slender,  terete.  Body  practically  destitute  of  pubescence  and 
setae.  Spiracular  disk  surrounded  by  three  very  broad  lobes,  a  flattened  ventral  lobe  and 
two  shorter  lateral  lobes  at  the  base  of  which  are  the  small  black  spiracles.  Gills  four, 
bluntly  rounded  and  developed  for  propulsion.  Head  capsule  of  four  elongate,  slender  rods 
or  plates,  interno-lateral  pair  forked  at  about  midlength.  Mandible  rather  small,  with  about 
three  blunt  lateral  teeth.  Antenna  elongate,  two-segmented. 

Pupa.- —  Cephalic  crest  setiferous.  Pronotal  breathing  horns  short,  blunt,  closely  applied 
to  thorax.  Mesonotum  precipitous,  at  crest  with  two  powerful  hooks  and  smaller  serrated 
plates  near  shoulder.  Wing  sheaths  reaching  end  of  second  abdominal  segment.  Leg 
sheaths  reaching  end  of  fourth  abdominal  segment,  middle  tarsi  the  shortest.  Abdomen 
with  a  transverse  row  of  setae  before  ends  of  segments. 

9 


946  CHARLES  PAUL  ALEXANDER 

Teucholabis  is  a  rather  extensive  genus  of  small  crane-flies  (including 
more  than  fifty  described  species)  which  find  their  center  of  distribution 
in  the  Tropics  of  the  New  World.  A  few  species  occur  in  Africa  and  the 
Oriental  region.  The  genotype,  Teucholabis  complexa,  is  the  only  species 
that  has  been  reared  (Johnson,  1900).  Johnson's,  material  was  kindly 
sent  to  the  writer  for  study,  and  furnishes  the  basis  for  the  following 
descriptions. 

Teucholabis  complexa  0.  S. 

1859     Teucholabis  complexa  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  223. 

Larvae  of  Teucholabis  complexa  were  found  by  Johnson  in  considerable 
numbers  beneath  the  bark  of  a  decayed  oak  below  Avalon,  New  Jersey, 
on  June  8,  1899.  They  commenced  pupating  about  the  13th,  the  imagines 
continuing  to  emerge  from  the  22d  to  the  27th.  This  gives  a  pupal 
duration  of  not  more  than  nine  days. 

Larva. —  Length,  9  mm. 

Diameter,  0.55-0.6  mm. 

Coloration  pale  yellowish  white,  spiracles  conspicuously  darker. 

Form  long  and  slender,  body  terete,  tapering  abruptly  to  the  small  prothoracic  segment 
(Plate  LXXIX,  424).  Sutures  between  segments  indistinct.  Body  practically  destitute  of 
pubescence  and  setae.  Spiracular  disk  (Plate  LXXIX,  427)  with  a  broad,  flattened,  ventral 
lobe,  which  is  very  bluntly  rounded  to  subtruncate  at  apex,  and  two  very  short,  blunt,  lateral 
lobes,  at  the  base  of  which  are  the  spiracles;  spiracular  disk  without  distinct  markings. 
Spiracles  small;  middle  piece  black,  ring  pale  horn-color;  spiracles  .rather  widely  separated, 
the  distance  between  them  about  equal  to  the  long  diameter  of  one.  Anal  gills  (Plate 
LXXIX,  428)  represented  by  four  blunt,  rounded  lobes,  which  are  apparently  developed  for 
propulsion  rather  than  for  a  respiratory  function. 

Head  not  easily  distinguishable  in  material  available  for  study.  Head  capsule  consisting 
of  four  long,  slender,  rodlike  plates,  the  internal  lateral  pair  forked  at  about  midlength, 
so  that  capsule  ends  in  six  rods.  Epipharynx  with  numerous  small  spines.  Antenna  (Plate 
LXXIX,  426)  two-segmented,  basal  segment  elongate-cylindrical,  apical  segment  small, 
ovate.  Mandible  (Plate  LXXIX,  425)  rather  small,  apical  point  inconspicuous,  with  about 
three  similar  lateral  teeth  below  it.  Lobes  of  the  maxilla  blunt,  stout,  hairy,  not  extending 
far  beyond  tip  of  mandible. 

Pupa. —  Length,  6.5-6.6  mm. 

Width,  d.-s.,  1-1.1  mm. 
Depth,  d.-v.,  1.2  mm. 

Coloration  pale;  head,  thorax,  and  sheaths  of  appendages  darker;  eyes  black. 
Form  slender,  narrowed  behind.     Between  antennal  bases  a  prominent,  two-parted  crest, 
each  lobe  somewhat  truncated  behind  and  bearing  a  single  stout  seta.     Front  above  eyes 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  947 

slightly  raised,  two  narrow  lines  on  front,  meeting  below  at  a  very  acute  angle.  Eyes  large 
in  male,  smaller  in  female,  the  front  correspondingly  narrowed  or  broadened.  Labrum  short, 
subtriangular.  Sheaths  of  labial  lobes  small,  suboval,  separated  by  tip  of  labrum.  Sheatha 
of  maxillary  palpi  short,  stout,  straight.  Antennal  sheaths  moderately  long,  extending  to 
just  beyond  wing  base;  basal  segments  prominent,  indicated  on  sheath  as  prominent  ele- 
vations. Pronotal  breathing  horns  short,  blunt,  anterior  face  closely  applied  to  pronotum, 
outer  face  free;  at  base  a  small  rounded  knob.  Thorax  very  deep,  precipitous,  at  crest 
(Plate  LXXIX,  429)  armed  with  two  strong  curved  hooks,  one  on  either  side  of  median  line; 
on  shoulder  laterad  of  these  hooks^  two  prominent  flattened  plates  whose  margins  are 
minutely  serrated,  the  dorsal,  or  outer,  plate  being  the  larger  and  terminating  in  a  large 
curved  hook.  Wing  sheaths  moderately  broad,  ending  just  before  tip  of  second  abdominal 
segment.  Leg  sheaths  (Plate  LXXIX,  430)  long  and  slender,  outer  pair  much  the  longest, 
ending  about  opposite  tip  of  fourth  abdominal  segment,  middle  pair  the  shortest. 
Mesonotum  with  two  strong  setae. 

Abdominal  segments  divided  into  two  annuli  by  an  indistinct  suture,  anterior  ring  narrow; 
anterior  ring  with  a  strong  seta  on  pleural  region;  posterior  ring  with  strong  setae  on  dorsum 
and  sternum  near  caudal  margin;  three  other  setae  on  pleural  region,  two  at  about  midlength 
of  segment,  the  third  near  base  and  more  dorsal  in  position.  Male  cauda  (Plate  LXXIX, 
431)  suddenly  narrowed,  terminating  in  two  blunt  ventral  lobes  and  two  acutely  pointed 
dorsal  lobes  bent  strongly  dorsad  at  their  tips  and  bearing  a  short  seta  in  notch  on  inner  face 
before  apex;  near  base  of  cauda,  on  dorsum,  a  broad  transverse  swelling  terminating  in  two 
widely  separated,  slender  tubercles,  immediately  cephalad  and  laterad  of  which  is  a  long  seta; 
three  strong  setae  on  either  side  near  base  of  cauda;  posterior  margin  of  segment  7  with  two 
powerful,  decussate  setae  on  dorsum,  and  between  them  two  small  setae;  another  powerful 
seta  near  pleural  region,  and  just  inside  still  another  smaller  seta. 

Nepionotype. —  Avalon,  New  Jersey,  June  8,  1899. 
Neanotype. —  With  type,  June  14,  1899. 
Paratypes. —  One  laf  va  and  one  pupa. 

Genus  Cladura  Osten  Sacken  (Gr.  branch  +  tail) 

1859    Cladura  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  229. 

The  genus  Cladura  includes  six  known  species  —  two  from  eastern  North 
America,  one  from  western  North  America,  and  three  from  Japan.  The 
small  Cladura  delicatula  Alex.,  of  the  mountainous  regions  of  the  north- 
eastern United  States,  differs  from  the  genotype,  C.  flavoferruginea  O.  S., 
in  several  important  respects  and  it  is  necessary  to  erect  a  new  group  to 
receive  it.  This  group  may,  for  the  present  at  least,  be  considered  as  a 
subgenus  of  Cladura,  and  the  name  Neocladura  (Gr.  new  +  Cladura)  is 
proposed.  Moreover,  the  genera  Crypteria  Bergr.  and  Pterochionea  Alex, 
are  closely  related  to  Cladura,  and  the  entire  group  are  almost  certainly  the 
direct  forbears  of  the  subapterous  genus  Chionea  Dalman.  Neocladura 


948  CHARLES  PAUL  ALEXANDER 

curiously  combines  the  structural  characters  of  Cladura,  Pterochionea, 
and  Crypteria.  In  the  long  basal  fusion-segment  of  the  antennal  flagellum 
and  in  the  wing  venation,  it  agrees  with  Pterochionea  and,  to  a  somewhat 
lesser  degree,  with  Cladura.  In  the  structure  of  the  male  hypopygium, 
which  has  two  slender  pleural  appendages,  it  departs  widely  from  the 
type  of  typical  Cladura,  Chionea,  and  Pterochionea,  and  agrees  better 
with  Crypteria,  as  well  as  with  Conosia  v.  d.  W.  and  Lecteria  O.  S. 
Because  of  this  combination  of  characters,  Neocladura  must  be  separated 
in  some  manner  from  Cladura  in  the  strict  sense.  It  must  be  borne  in 
mind,  however,  that  the  conspicuous  difference  in  the  structure  of  the 
male  hypopygium  probably  has  a  phylogenetic  significance,  and  the  two 
flies  may  not  be  so  closely  united  as  this  arrangement  would  indicate.^ 
The  habits  of  the  adult  flies  of  Cladura  and  Neocladura  are  generally 
similar.  Both  species  fly  in  late  summer  and  in  the  autumn.  They 
often  occur  on  dry,  wooded  hillsides  remote  from  streams  and  other 
bodies  of  water.  The  habits  of  the  adult  flies  of  Cladura  flavoferruginea 
have  been  discussed  by  the  writer  in  an  earlier  paper  (Alexander,  1910:250, 
as  C.  indivisa).  His  observations,  made  near  Gloversville,  New  York,  on 
September  22,  1909,  are  as  follows: 

I  went  to  Simmon's  Woods,  southeast  of  Gloversville,  New  York,  this  afternoon,  and 
was  very  agreeably  surprised  at  the  occurrence,  in  large  numbers,  of  this  usually  uncommon 
insect.  Near  the  entrance  of  the  woods,  where  Simmon's  Brook  emerges,  the  insects  were 
found  in  numbers.  At  each  step  they  flew  out  of  the  bushes  to  others  farther  away.  They 
are  wary  insects,  and  when  sitting  on  the  upper  side  of  a  leaf,  slip  over  the  edge  and  hang 
inverted  from  the  lower  side  when  alarmed  by  an  observer.  They  present  a  very  character- 
istic attitude,  sitting  on  the  leaf  of  a  tree,  with  their  wings  folded  flat  over  the  abdomen,  and 
the  six  long  legs  stretched  out  over  the  leaf.  A  few  were  taken  in  copulation;  these  were 
all  hanging  on  the  under  side  of  a  leaf.  Their  habit  of  clinging  to  the  under  surface  of  a  leaf 
is  quite  remarkable  and  I  found  several  by  looking  for  them  there. 

There  were  hundreds  of  specimens  in  the  low  bushes  of  the  woods,  usually  on  the  broad 
leaves  of  deciduous  trees  at  a  height  of  two  or  three  feet.  Sometimes  they  would  alight 
on  hemlock,  and,  occasionally,  in  ferns  near  the  ground.  It  was  the  only  tipulid  observed 
here  to-day. 

The  eggs  of  C.  delicatula  have  been  taken  from  gravid  females.  They 
are  comparatively  few  in  number,  but  because  of  their  unusual  size  they 
almost  fill  the  entire  abdominal  cavity  of  the  fly.  These  large  eggs  are 
elongate-ovate  in  shape. 

Cladura  flavoferruginea  O.  S. 

1859    Cladura  flavoferruginea  0.  S.      Proc.  Acad.  Nat.  Sci.  Phila.,  pi.  4,  fig.  34. 
1861     Cladura  indivisa  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  291. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  949 

The  immature  stages  of  Cladura  flavoferruginea  were  discovered  while 
this  paper  was  in  press.  A  brief  account  of  the  larvae  and  pupae  are 
given  here  in  order  to  complete  the  data. 

The  larvae  were  found  in  Augurville  Woods  near  Urbana,  Illinois. 
They  occurred  in  soil  which  was  baked  hard  and  dry  and  which  supported 
scarcely  any  other  insect  life.  Associated  with  the  larvae  when  first 
discovered  were  larvae  of  a  scarabaeid,  Xyloryctes  satyrus;  a  tenebrionid, 
Meracantha  contracta',  a  few  dipterous  larvae  of  the  genera  Sciara  and  Psilo- 
cephala;  millepedes  of  the  genus  Spirobolus;  and  a  few  less  common  forms 
of  animal  life.  A  layer  of  dead  leaves  and  other  decaying  vegetable  matter 
covered  the  surface,  but  this  had  not  prevented  an  almost  complete  drying 
out  of  the  soil  to  a  depth  varying  from  six  inches  to  more  than  a  foot. 
The  only  other  tipulid  larvae  characteristic  of  such  dry  soil  are  species  of 
Dicranoptycha  (page  828). 

The  most  conspicuous  features  of  the  pupa  are  its  exceedingly  small 
size  as  compared  with  the  adult  fly  that  emerges  from  it,  and  the  entire 
lack  of  protuberant  pronotal  breathing  horns. 

Larva. —  Length,  10-10.5  mm. 
Diameter,  1.2  mm. 

Coloration  light  yellow  thruout. 

Form  comparatively  short  and  stout.  Integument  provided  with  a  delicate  appressed 
pubescence;  no  distinct  setae.  Basal  annulus  of  abdominal  segments  2  to  7  with  a  transverse 
area  of  microscopic  points  arranged  in  long  transverse  rows;  last  ventral  segment  with  a  flat- 
tened lobe  covered  with  short  setae,  evidently  an  organ  for  shoving.  Spiracular  disk  entirely 
without  lobes,  the  spiracles  being  located  on  the  exposed  dorso-caudal  surface  of  the  last 
abdominal  segment.  Head  capsule  relatively  compact;  frontal  plate  broad,  only  slightly 
narrowed  behind.  Labrum  quadrate,  with  conspicuous  oval  lateral  arms.  Antenna  two- 
segmented,  terminal  segment  elongate-oval.  Mandibles  of  a  herbivorous  type,  with  an  apical 
point  and  two  incomplete  rows  of  teeth  on  inner,  or  cutting,  face.  Mental  bars  widely 
separated,  each  bar  provided  with  two  acute  teeth  at  its  proximal  end. 

Pupa. —  Length,  6.7  mm. 
Width,  1.4  mm. 
Depth,  1.4  mm. 

Coloration  pale  yellow;  head,  thorax,  and  appendages  darkening  in  age. 

Cephalic  crest  gibbous,  entire  or  feebly  bifid,  armed  on  either  side  with  a  single  powerful 
bristle;  two  bristles  on  vertex  and  two  on  front;  labrum  with  a  pair  of  small  bristles  at  each 
cephalic  lateral  angle.  Labial  lobes  subquadrate,  weakly  separated  by  apex  of  labral  sheath. 
Palpal  sheaths  short  and  stout,  straight.  Lateral  margin  of  eye  produced  laterad  into  a 
digitiform  lobe.  Antennal  sheaths  extending  to  opposite  one-third  length  of  wing  sheaths. 
Pronotal  breathing  pores  entirely  sessile.  Pronotum  and  mesonotum  armed  with 


950  CHARLES  PAUL  ALEXANDER 

conspicuous  bristles.  Wing  sheaths  ending  opposite  base  of  third  abdominal  segment. 
Leg  sheaths  unusually  long,  ending  opposite  base  of  sixth  abdominal  segment;  hind  legs  the 
longest,  middle  legs  the  shortest.  Abdominal  tergites  with  ten  strong  bristles,  eight  being 
arranged  in  a  single  transverse  row  along  posterior  margin;  abdominal  pleurites  with  four 
strong  bristles,  one  on  each  anterior  ring,  two  near  caudal  margin  of  posterior  ring,  one 
ventrad  of  spiracle,  the  last-named  rudimentary,  located  on  segments  2  to  7;  sternites  unarmed 
with  bristles. 

Genus  Chionea  Dalman  (Gr'snow) 

1816     Chionea  Dalm.     K.  Vet.  Akad.  Handl.,  vol.  1,  p.  102. 

1912    Sphaconophilus  Beck.    Ann.  Soc.  Ent.  Belgique,  vol.  56,  p.  142. 

Chionea  is  a  small  genus  of  nearly  apterous  crane-flies,  found  thruout 
the  North  Temperate  Zone.  All  of  the  known  species  (about  eight) 
have  the  wings  reduced  to  mere  vestiges,  this  being  the  only  genus  of 
considerable  size  in  which  all  the  species  show  this  condition.  The  South 
African  genus  Platylimnobia  Alex.,  which  shows  a  somewhat  similai 
condition  of  wing  atrophy,  is  probably  not  very  closely  allied  to  Chionea. 

The  adult  flies  of  Chionea  are  most  often  found  walking  awkwardly 
over  the  snow  in  the  late  fall  or  early  spring  or  during  warm  days  in 
winter.  A  few  instances,  however,  have  been  recorded  in  which  specimens 
were  found  with  the  temperature  below  freezing.  At  other  seasons  of 
the  year  they  may  be  found  among  fallen  leaves,  under  moss  and  stones, 
in  the  nests  of  small  mammals  as  Arvicola  (Schmitz,  1914),  or  in  deserted 
subterranean  wasps'  nests  (Schmitz,  1916,  and  Becker,  1912).  Interesting 
accounts  of  the  habits  of  the  adult  flies  of  the  commonest  local  species, 
Chionea  valga  Harris,  are  supplied  by  Ainslie  (1906),  Johnson  (1907), 
Washburn  (1907),  and  others.  Lugger  (1896)  gives  the  following  interest- 
ing account  of  the  habits  of  the  same  species: 

As  a  general  rule  the  wingless  flies  are  found  only  early  in  the  morning,  though  in  one  case 
a  female  was  discovered  crawling  over  the  snow  in  the  evening.  ...  A  few  winters  ago 
the  writer  discovered  a  female  moving  slowly  over  the  snow  and  by  searching  he  soon  detected 
a  male.  Putting  both  together  under  an  inverted  glass  the  snow-flies  immediately  mated, 
notwithstanding  it  was  several  degrees  below  the  freezing  point.  Soon  afterward  the  female 
found  a  crack  in  the  glassy  surface  of  the  frozen  enow,  and  forcing  herself  into  it  slowly 
disappeared  from  view.  Penetrating  for  some  depth  into  the  snow  she  deposited  a  number 
of  elongated  eggs,  which,  however,  did  not  hatch.  Most  females  found  seemed  to  have  the 
eggs  already  fully  matured  and  only  lacked  to  be  fertilized. 

Frauenfeld  (Brauer,  Egger,  and  Frauenfeld,  1854:616)  believed  that 
the  eggs  are  deposited  in  snow,  since  they  are  often  laid  in  January  or 
February.  He  thought  the  slimy  substance  secreted  by  the  small  lateral 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  951 

vesicles  of  the  receptaculum  seminis  of  the  female  acts  as  a  covering  to 
protect  the  eggs  from  wet  and  cold.  It  seems  probable,  however,  that 
the  flies  enter  some  crevice  in  the  snow  around  the  base  of  trees  or  shrubs 
and  reach  solid  earth,  at  least  in  some  cases.  Many  authors  have  held 
the  flies  to  be  nocturnal  in  their  habits.  Recently,  Marchand  (1917) 
has  furnished  some  interesting  notes  on  an  alpine  Chionea,  presumably 
C.  alpina  Bezzi.  His  observations  and  experiments  showed  him  that 
Chionea  was  perfectly  adapted  to  life  on  the  snow,  being  attracted  to 
this  medium  by  its  bright  light  and  white  color,  the  contact  of  the  cold 
surface  on  the  feet  resulting  in  a  direct  stimulus  thru  the  claws.  The 
insects  drink  water  by  pressing  their  proboscides  against  the  snow.  Mar- 
chand considers  the  principal  reason  for  these  activities'  Ibeing  held  on 
the  snow  to  be  for  the  purpose  of  mating,  since  the  insects  can  cover 
considerable  distances  over  the  level  surfaces  and  are  much  more  visible 
to  one  another  at  this  time.  The  copulation  of  this  crane-fly  has  been 
fully  described  by  Mik  (Osten  Sacken,  1887:196)  as  follows: 

The  upper  valves  of  the  ovipositor  prevent  the  male  from  getting  on  the  back  of  the  female; 
it  lies  on  its  own  back,  in  the  direction  of  the  longitudinal  axis  of  the  body  of  the  female; 
when  the  latter  is  walking  it  drags  the  male,  who  raises  himself  on  his  hind  legs  to  an  almost 
perpendicular  position;  this  serves  to  explain  the  unusual  incrassation  cf  these  legs. 

The  genotype,  C.  araneoides  Dalm.,  is  the  only  species  whose  immature 
stages  have  been  described. 

Chionea  araneoides  Dalm. 

1816  Chionea  araneoides  Dalm.  K.  Vet.  Akad.  HandL,  vol.  1,  p.  104/ 
The  present  knowledge  of  the  life  history  of  Chionea  araneoides  is  due 
almost  entirely  to  the  work  of  Brauer,  Egger,  and  Frauenfeld  (1854). 
In  February  these  investigators  brought  living  adults,  taken  in  copula, 
into  an  unheated  room  and  placed  them  with  their  natural  surroundings, 
such  as  rich,  damp,  humous  earth,  rotten  grape  leaves,  and  similar  sub- 
stances, in  glass  containers.  After  a  time  a  great  number  of  small, 
elongated  eggs  of  a  hyaline  appearance  were  noted,  laid  at  random,  some 
being  deposited  on  the  walls  of  the  container,  where  they  adhered  but 
soon  shrunk  and  appeared  dried  out.  Unfortunately  the  duration  of  the 
egg  stage  was  not  ascertained.  Some  weeks  later  the  young  larvae  were 
found  in  the  soil.  They  were  of  a  cylindrical  form,  very  pale  yellow  in 
color  and  not  especially  active.  They  were  associated  with  numerous 


952  CHARLES  PAUL  ALEXANDER 

larvae  of  Sdara  longipes  Meig.  Toward  the  end  of  May  only  a  few  larvae 
were  left,  and  these  appeared  fully  grown  but  were  not  carried  over  into 
the  pupal  condition,  which  is  still  unknown. 

Larva.- —  Length  when  fully  grown,  7.4  mm. 

Color  light  reddish  yellow,  contents  of  alimentary  tract  showing  thru  body.  Body  terete 
(Plate  LXXXII,  443),  consisting  of  twelve  segments,  there  being,  besides  the  head,  three 
thoracic  and  nine  abdominal  segments;  last  segment  wedge-shaped  (Plate  LXXXII,  446 
and  447),  obliquely  truncated  so  that  the  surface  slopes  from  behind  upward.  Two  spir- 
acles on  this  oblique  surface,  with  a  paler  brown,  pincer-shaped  mark  between.  Mouth 
parts  powerfully  constructed.  Mandible  (Plate  LXXXII,  444),  produced  into  a  long  apical 
point;  near  midlength  on  inner  edge  of  mandible  a  deep  incision,  distad  of  this  about  five 
teeth,  basad  of  it  three  teeth.  Labrum  quadrangular,  with  two  lateral  points  on  anterior  part. 
Men  turn  projecting  outward  as  a  cuticular  rounded  lobe.  (Brauer  shows  two  other  append- 
ages which  are  toothed  on  the  outer  face  [Plate  LXXXII,  445,  of  this  paper] ;  these  probably 
represent  part  of  the  mentum,  but  from  Brauer's  figures  they  would  appear  to  lie  above 
the  labrum.) 

Subtribe  Elephantomyaria 

Genus  Elephantomyia  Osten  Sacken  (Gr.  elephant  +  fly) 

1859     Elephantomyia  O.  S.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  220. 

Larva.—  Form  terete.  Segments  of  body  just  before  sutures  with  transverse  rows  of  stiff 
hairs;  abdominal  segments  5  to  8  on  ventral  surface  with  a  mouthlike  depression  surrounded 
by  long,  stiff  hairs.  Spiracular  disk  surrounded  by  four  lobes,  ventral  pair  the  longest, 
each  of  the  latter  bearing  at  its  tip  a  single  very  long  bristle.  Spiracles  moderately  large, 
located  at  base  of  lateral  lobes.  Head  capsule  very  long  and  narrow,  the  four  plates  very 
elongated.  Mandible  very  small.  Maxillary  palpi  short.  Esophageal  region  surrounded 
by  chitinized  plates  which  are  conspicuously  obliquely  ridged.  Body  of  larva  covered  with 
numerous  long,  appressed  hairs,  producing  a  satiny  appearance.  Coloration  saturated 
golden  yellow. 

Pupa.—  Form  slender.  A  small  crest  on  vertex  above  eyes.  Eyes  very  large,  globular, 
narrowly  separated  on  frontal  and  vertical  regions,  ilostral  sheath  very  elongated;  palpi 
recurved  against  it.  Antennal  sheaths  lying  across  eye.  Head  and  thorax  with  setiferous 
tubercles.  Abdomen  with  rudimentary  lateral  spiracles. 

Elephantomyia  is  a  small  genus  of  crane-flies  including  about  a  dozen 
described  living  species.  These  are  found  in  widely  separated  regions 
of  the  world,  there  being  about  four  in  the  East  Indies,  five  in  Africa, 
and  two  in  North  America,  one  of  which  has  been  recorded  also  from 
Europe.  The  genus  is  found  fossil  in  the  Baltic  amber.  The  adult 
flies  are  remarkable  for  their  very  elongated  rostra,  and  undoubtedly 
they  feed  on  the  nectar  of  tubular  flowers  as  in  the  related  genus  Toxor- 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  953 

hina.  No  records  are  available,  however,  to  indicate  what  species  of 
plants  are  thus  frequented.  The  larvae  live  in  decaying  wood.  The 
only  species  whose  immature  stages  are  known  is  the  genotype,  Elephanto- 
myia  westwoodi.  The  literature  on  the  immature  stages  of  this  group 
of  flies  is  very  limited,  the  only  record  being  the  unknown  Limnobiine 
No.  1  of  Malloch  (19 15-17  b:  235-236),  who  gives  a  good  description  of 
the  larva.  The  pupa  is  here  described  and  figured  for  the  first  time. 

Elephantomyia  westwoodi  O.  S. 

1869     Elephantomyia  westwoodi  O.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  109. 

Elephantomyia  westwoodi  is  a  rather  common  fly  in  eastern  North 
America.  The  adults  are  usually  found  in  cool,  shaded  woods  and  may 
be  swept  from  vegetation  in  these  localities.  The  larva  lives  in  decaying 
wood,  as  is  shown  by  the  three  records  available  to  the  writer. 

The  larvae  were  found  by  Shannon  in  a  very  wet,  rotten,  willow  log 
lying  near  the  tidal  flat  above  the  brewery  at  Rosslyn,  Virginia,  on  May 
21,  1913.  About  eight  larvae  were  found,  well  scattered  thru  the  log. 
They  were  very  active  in  their  movements  and  were  very  beautiful,  in 
life  being  of  a  deep  golden  yellow  color.  These  larvae  were  placed  in 
rearing,  and  adults  issued  on  May  27  and  29  and  June  5  and  7,  indicating 
a  pupal  duration  of  a  week  or  slightly  less.  Larvae  were  found  in  this 
log  also  on  November  23,  1912,  and  at  that  time  they  were  almost  grown, 
being  about  one-half  inch  in  length.  This  shows  that  the  species  spends 
the  winter  as  almost  fully  grown  larvae. 

Johnson  found  these  larvae  near  Edge  Hill,  Pennsylvania,  on  June 
25,  1899,  and  on  May  25,  1905,  in  a  log,  just  beneath  the  bark.  Larvae 
and  pupae  obtained  by  him  were  in  the  material  studied  in  the  preparation 
of  this  paper. 

Malloch  found  a  single  larva  of  this  species  in  a  much  decayed  log  at 
White  Heath,  Illinois,  on  April  30,  1916  (Malloch,  1915-17 b: 236). 

Larva.—  Length,  10-13.4  mm. 
Diameter,  1.2-1.3  mm. 

Coloration  of  body,  a  deep  saturated  golden  yellow  thruout. 

Body  moderately  elongated,  terete,  relatively  slender,  tapering  gradually  toward  either  end 
but  more  noticeably  and  abruptly  toward  anterior  end  (Plate  LXXX,  432) ;  the  three  thoracic 
segments  gradually  increasing  in  size  from  the  prothoracic  backward.  Abdominal  segments 
1  and  2  short,  the  third  to  the  fifth  the  longest,  remaining  segments  gradually  shorter.  Sur- 


954  CHARLES  PAUL  ALEXANDER 

face  of  body  densely  covered  with  long,  satiny,  appressed  hairs;  thoracic  and  abdominal 
segments  with  a  dorsal  and  a  ventral  transverse  ridge  of  short,  stiff  hairs  just  before  caudal 
margins  of  segments,  these  being  longest  on  lateral  parts  of  ridge;  dorsal  segments  6  and 
7  with  incomplete  transverse  rows  at  about  rnid length;  on  ventral  surface  of  abdominal  seg- 
ments 1  to  3,  near  base,  two  transverse  rows  of  tiny  spines  with  a  depressed  area  between; 
segment  4  without  this  distinct  double  ridge;  segments  5  to  7  with  a  very  conspicuous 
mouthlike  depression  at  base  of  each,  with  a  liplike  margin  on  either  side,  the  anterior  margin 
narrow  with  stiff  hairs,  the  posterior  margin  swollen  and  densely  set  with  tiny  spines;  on  seg- 
ment 8  the  transverse  mouth  lying  near  end  of  segment,  its  anterior  lip  with  a  dense  fringe 
of  long  hairs  directed  backward.  Spiracular  disk  (Plate  LXXX,  434)  surrounded  by  four 
lobes;  lateral  lobes  the  shortest,  blunt  at  tips,  inner  face  of  each  lobe  slightly  expanded 
at  tip,  a  few  short  silky  hairs  on  outer  face  of  lobe,  inner  face  slightly  chitinized;  ventral  lobes 
longer,  broad,  tapering  gradually  to  obtuse  tips,  notch  between  lobes  deep,  V-shaped,  a 
narrow  fringe  of  short,  dense,  golden,  recurved  hairs  along  outer  face;  at  tip  of  each  ventral 
lobe  a  single  long,  powerful  bristle.  Spiracular  disk  unmarked.  Spiracles  large,  very 
widely  separated,  situated  at  base  of  lateral  lobes. 

Head  capsule  (Plate  LXXX,  433)  very  small  and  narrow,  the  plates,  four"  in  number, 
being  greatly  elongated.  (The  exact  details  are  difficult  to  see  in  the  material  available  for 
study.)  Labrum  broad,  anterior  margin  evenly  rounded,  with  a  few  long  hairs.  Epipharyn  • 
geal  region  provided  with  long  setae  directed  backward.  Mandible  very  small,  base 
narrowed,  tip  produced  into  an  acute  point  with  smaller  teeth  at  about  midlength.  Mental 
region  feebly  chitinized.  Hypopharynx  semicircular,  rounded,  anterior  margin  delicately 
grooved.  Esophageal  region  elongated,  inclosed  by  chitinized  plates  provided  with  parallel 
ridges  running  outward  on  one  side  and  inward  on  opposite  face,  upon  focusing  produc- 
ing a  latticed  appearance.  Maxilla  densely  hairy.  Antenna  two-segmented,  conspicuous; 
basal  segment  short,  apical  segment  larger,  suboval.  Sides  of  capsule  on  swelling  behind 
the  maxilla  with  a  brush  of  very  long  hairs.  Plates  of  capsule  elongated,  expanded 
and  chitinized  at  tips  and  along  margins;  ventral  plate  near  esophageal  region  with 
setae. 

Pupa. —  Length,  8.4  mm. 

Width,  d.-s.,  1.1  mm. 
Depth,  d.-v.,  1.3  mm. 

Coloration  light  yellowish  brown;  abdomen  darker;  thoracic  dorsum,  sheaths  of  legs, 
and  ovipositor  more  yellowish.  (In  younger  pupae  the  coloration  is  more  uniformly 
pale.) 

Form  slender,  body  narrow.  Eyes  very  large,  rounded  (Plate  LXXXI,  441).  Antennal 
sheaths  lying  directly  across  face  of  eye.  Vertex  with  a  small  but  prominent  crest  lying 
transversely  above  eyes;  cephalad  of  this  crest  and  just  proximad  of  the  antenna!  bases,  a 
setiferous  tubercle.  Front  between  eyes  moderately  broad,  produced  caudad  into  the  very 
elongate  rostral  sheath,  which  is  transversely  wrinkJ3d.  Sheaths  of  maxillary  palpi  recurved, 
lying  alongside  rostrum.  Sheaths  of  labium  tiny,  bilobed,  lying  at  tip  of  rostrum.  Two 
long  setae  on  front  between  eyes.  Eyes  very  large,  narrowly  separated  on  dorsum  of  head, 
the  hinder  part  sunken  under  pronotum.  Two  conspicuous  black  tubercles  behind  antennal 
bases,  each  bearing  a  long,  stout  seta.  Pronotal  breathing  horns  (Plate  LXXX,  435)  small, 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  955 

short,  clavate,  yellow,  constricted  at  bases,  which  are  blackened;  a  prominent  seta  just  above 
each  breathing  horn;  two  setae  on  each  side  of  pronotum  beneath  eye.  Mesonotal  prescutum 
strongly  convex,  with  six  strong  setae  on  each  side,  one  just  cephalad  of  base  of  wing,  another 
at  joint  of  wing,  and  two  semitransverse  groups  in  front  of  and  behind  level  of  wing  base. 
Wing  sheaths  short,  ending  some  distance  beyond  tips  of  hind  tibiae  and  just  beyond  base 
of  third  abdominal  segment.  Leg  sheaths  (Plate  LXXXI,  442)  with  fore  femora  strongly 
swollen,  lying  alongside  rostral  sheath;  tips  of  middle  tibiae  ending  just  beyond  tips  of  fore 
tibiae;  legs  very  long,  ending  just  before  tip  of  fifth  abdominal  segment. 

Abdominal  segments  densely  and  microscopically  punctulate.  Segments  2  to  7  with  two 
narrow  basal  rings  and  a  broad  posterior  ring.  Setae  of  dorsal  abdominal  segments  (Plate 
LXXX,  437)  en  the  posterior  ring  consisting  of  a  caudal  series  of  two  strong  outer  setae  and 
two  smaller  inner  ones,  the  outer  one  of  the  inner  series  close  to  the  proximal  one  of  the  outer 
series;  lateral  series  of  setae  powerful;  basal  series  in  alinement  with  anterior  lateral  seta, 
consisting  of  two  powerful  outer  setae  and  a  delicate  inner  one;  pleural  segments  with  a  rudi- 
mentary spiracle  on  segments  2  to  7,  each  spiracle  with  a  stout  seta  above  it;  setae  of  ventral 
segments  (Plate  LXXX,  436)  with  two  strong  bristles  on  each  side  near  posterior  lateral  mar- 
gin, the  outermost  with  a  tiny  seta  above  it;  a  single  basal  seta  located  on  a  level  with  ves- 
tigial spiracles.  Female  cauda  (Plate  LXXX,  439)  with  sternum  of  eighth  segment  having 
two  sharp,  curved,  widely  separated  spines,  and  just  laterad  of  these  a  stout  seta;  pleural 
region  with  another  seta  on  same  level;  sternal  valves  of  ovipositor  only  a  little  shorter  than 
tergal  valves;  dorsum  of  segment  8  with  two  long  teeth,  above  which  are  two  slender,  diver- 
gent tubercles;  tergal  valves  broad  basally,  narrowed  suddenly  at  tip,  on  either  side  with  a 
small,  subapical  seta.  Male  cauda  (Plate  LXXX,  438  and  440)  with  eighth  sternite  hav- 
ing a  strong  median  tubercle  that  is  two- toothed;  laterad  of  this  a  strong  chitinized  tooth 
bearing  a  powerful  seta  on  outer  face;  a  similar  strong  lateral  seta;  eighth  tergite  with  four 
strong  tubercles  arranged  to  form  a  square;  ninth  sternite  blunt,  each  lobe  ending  in  two 
small  tubercles;  ninth  tergite  ending  in  two  divergent  lobes  bearing  at  tip  a  large  and  a  small 
seta  and  on  dorsal  face  at  about  midlength  another  strong  seta. 

Nepionotype. —  Rosslyn,  Virginia,  May  21,  1913. 

Neanotype. —  Edge  Hill,  Pennsylvania,  May  25,  1905. 

Paratypes. —  Larvae  with  type  larva  (two)  and  with  type  pupa  (two). 

Genus  Toxorhina  Loew  (Gr.  bow  +  nose) 

1835  Limnobiorhynchus  Westw.     Ann.  Soc.  Ent.  France,  p.  683  (spurious  name). 

1851  Toxorhina  Loew.     Linnaea  Entomol.,  vol.  5,  p.  400. 

1869  Toxorrhina  O.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  109-114. 

1910  Neoceratocheilus  Wesch6.     Journ.  Linn.  Soc.,  ZooL,  vol.  30,  p.  358. 

Toxorhina  is  a  small  genus  including  about  nine  described  species, 
almost  all  of  which  are  from  the  New  World.  Two  species  occur  in 
Africa  and  one  in  India.  Toxorhina  madagascariensis  Meun.  is  described 
from  African  copal  (Pleistocene).  Nothing  has  been  published  con- 
cerning the  immature  stages  of  any  member  of  this  genus. 


956  CHARLES  PAUL  ALEXANDER 

Toxorhina  muliebris  (O.  S.) 

1865     Tox&rrhina  muliebris  0.  S.     Proc.  Ent.  Soc.  Phila.,  p.  233. 

Toxorhina  muliebris  is  the  commonest  species  of  the  genus  in  the 
United  States,  with  a  rather  wide  range  thruout  the  Northeastern  States. 
The  adult  flies  suck  nectar  from  various  flowers,  such  as  the  following: 
Rhamnaceae,  Ceanothus  americanus  Linn.  (Banks);  Ericaceae,  Clethra 
alnifolia  Linn.  (McAtee);  Apocynaceae,  Apocynum  medium  Greene 
(McAtee);  Compositae,  Solidago  canadensis  Linn.  (Knab). 

The  immature  stages  are  unknown,  but  from  Mrs.  Tothill's  tent-trap 
observations  they  are  presumably  spent  in  mud,  since  adult  flies  were 
found  in  her  traps  set  over  wet,  sedgy  spots  near  Ithaca,  New  York.  It 
may  be,  however,  that  the  insects  live  in  fragments  of  decaying  wood 
which  might  be  buried  in  this  mud,  since  such  a  habitat  conforms  more 
closely  to  that  of  Elephantomyia,  which  is  apparently  closely  related  to 
Toxorhina. 

Eriopterine  No.  1 

A  very  curious  larva,  which  has  not  been  reared,  has  been  found  in 
various  places  near  Ithaca  during  the  past  few  years.  It  is  a  small,  pale 
larva,  very  delicate  and  almost  diaphanous  in  appearance,  at  the  posterior 
end  with  five  flattened  black  plates  with  serrated  margins,  and  with'  its 
thoracic  segments  capable  of  considerable  lateral  extension.  The  larva 
is  undoubtedly  'an  eriopterine,  but  it  introduces  a  type  of  spiracular 
disk  that  has  not  been  found  elsewhere  in  the  tribe.  The  writer  finds 
it  difficult  to  believe  that  this  curious  larva  can  belong  to  any  of  the 
eriopterine  genera  discussed  in  this  paper,  and  yet  there  are  very  few 
possibilities  remaining;  and  one  of  these  (Cryptolabis)  does  not  occur 
in  the  habitat  frequented  by  this  larva.  The  genus  Atarba,  whose 
immature  stages  are  still  wholly  unknown,  is  a  possibility.  Empeda,  which 
the  writer  considers  to  be  a  subgenus  of  Erioptera,  has  not  been 
reared  and  must  also  be  considered  as  a  possibility.  If  this  is  the  larva 
of  Empeda,  the  group  at  once  assumes  full  generic  rank  as  given  it  by 
Osten  Sacken,  but  occupying  ah  isolated  position  and  no  closer  to  Gono- 
myia  than  to  Erioptera.  The  larvae  of  this  species  were  found  commonly 
on  Bool's  hillside,  at  Ithaca,  where  they  occurred  in  association  with 
numerous  other  crane-fly  larvae  discussed  elsewhere  (page  781).  The 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  957 

larva  is  described  here  in  the  hope  that  it  will  be  reared  and  its  identity 
ascertained. 

Larva. —  Length,  7-8.2  mm. 

Diameter,  0.4-0.5  mm. 

Coloration  pale  yellowish  white;  skin  very  delicate,  almost  diaphanous. 

Body  moderately  elongated,  terete;  meso-  and  metathoracic  segments  (Plate  LXXV,  404) 
and  eighth  abdominal  segment  capable  of  considerable  expansion  laterally,  and,  in  death, 
usually  greatly  swollen;  last  segment  of  body  narrowed,  cylindrical,  with  a  number  of  long 
setae,  including  a  group  of  five  near  base  of  lateral  lobes.  A  few  scattered  setae  along  abdom- 
inal segments.  Spiracular  disk  (Plate  LXXV,  405)  surrounded  by  five  equal  elongate  spatu- 
late  blades,  these  blades  flattened,  jet-black  in  color,  margins  finely  toothed;  paired  lobes 
near  base  with  a  subhy aline  median  spot;  margins  of  lobes* (Plate  LXXV,  406)  with  twenty- 
five  to  thirty  hooks,  recurved  ones  alternating  with  others  laterally  directed;  when  blades  are 
closed,  these  margins  hooking  closely  together;  at  ends  of  blades  and  sparsely  scattered  along 
margin,  long,  delicate  setae;  at  apex  of  blades,  two  bristles;  no  spiracles  found  at  base  of 
lobes.  Anal  gills  four,  lateral  pair  elongated,  telescopic,  inner  pair  shorter. 

Head  capsule  much  as  in  other  eriopterine  genera,  especially  Ormosia  and  Gonomyia, 
dorsal  plates  slender,  ventral  bars  a  little  longer.  Labrum  as  in  the  tribe;  epipharyngeal 
region  with  a  large  apical  setiferous  pad  and  two  smaller  pads  nearer  base.  Mentum  not 
formed  of  ventral  bars  of  capsule  as  in  Molophilus.  Hypopharynx  a  semicircular  cushion 
covered  with  long,  dense  setae,  their  tips  a  little  recurved.  Antenna  with  apical  papilla 
very  long  for  this  tribe,  about  equal  in  length  to  basal  segment,  cylindrical,  with  tip  rounded. 
Mandible  moderately  large,  apical  tooth  not  prominent,  lateral  teeth  rather  conspicuous, 
basal  ones  smaller  but  not  so  excessively  reduced  as  in  other  members  of  the  tribe;  prostheca 
large. 

(Described  from  larvae,  Ithaca,  New  York,  May  11,  1917.    No.  29-1917.) 

Tribe  Styringomyiini 

The  Styringomyiini  comprise  a  small  group  of  very  peculiar  crane-flies 
with  a  tropicopolitan  distribution.  There  is  only  the  single  genus, 
Styringomyia,  with  about  twenty-five  described  species.  Most  of  the 
species  are  from  tropical  Africa  and  Asia,  tho  a  few  range  into  Australia 
and  the  Hawaiian  Islands,  and  one,  Styringomyia  americana  Alex.,  is 
found  in  tropical  South  America. 

Genus  Styringomyia  Loew  (Gr.  a  kind  of  tree-gum  -{-fly) 

1845    Styringomyia  Loew.     Dipt.  Beitr.,  vol.  1,  p.  6.     (Correctly  Syringomyia  — 

roth  in  litt.) 

1903    Idiophlebia  Griinb.     Zool.  Anzeig.,  vol.  26,  p.  524-528. 
1912     Pycnocrepis  Enderl.     Zool.  Jahrb.,  vol.  32,  part  1,  p.  65. 
1917    Mesomyites  Ckll.     Proc.  U.  S.  Nat.  Mus.,  vol.  52,  p.  377. 


958  CHARLES  PAUL  ALEXANDER 

The  history  of  the  genus  Styringomyia  is  remarkable.  It  was  erected 
by  Loew  in  1845,  being  based  on  the  fossil  species  Styringomyia  venusta 
Loew,  from  African  copal.  Many  years  later  it  was  found  to  be  still 
living  in  the*  Tropics  of  both  hemispheres.  The  earliest  fossil  records 
pertain  to  the  Oligocene  of  northern  Europe  (page  765). 

The  adult  flies  have  such  a  curious  structure  that  it  seems  best  to 
remove  them  from  the  former  tribe  Antochini,  where  they  have  long  been 
placed.  Concerning  the  first  living  species  to  be  described,  the  Hawaiian 
S.  didyma  Grimsh.,  Perkins  (1913 : clxxxii)  says: 

It  sometimes  swarms  at  night  around  the  electric  lights,  sitting  quietly  on  the  walls  and 
ceilings,  with  the  body  pressed  closely  to  the  surface,  and  the  front  and  middle  legs  extended 
straight  forward  in  front  of  the  head  in  a  characteristic  manner. 

Annandale  has  made  similar  observations  on  the  resting  positions  of  S. 
ceylonica  Edw.,  taken  in  India.  He  says  (cited  by  Edwards,  1914-15 : 207) : 
"  This  species  rests  on  walls  with  the  two  anterior  pairs  of  legs  stretched 
out  straight  in  front  and  the  posterior  pair  behind,  resembling  a  stray 
piece  of  cobweb."  Jacobson  has  recorded  much  the  same  for  S.  jacobsoni 
Edw.  (De  Meijere,  1911:41-42,  as  S.  didyma)  in  Java.  The  flies  are 
attracted  to  lamps  and  are  almost  always  to  be  found  in  copulation,  the 
head  of  one  directed  away  from  the  other;  while  thus  engaged,  some- 
times one,  sometimes  the  other,  will  run  forward  for  a  short  distance, 
producing  a  peculiar  appearance.  Munro  (in  litt.)  reports  that  the  habits 
of  S.  vittata  Edw.  as  observed  in  eastern  Transvaal  in  late  April,  1920, 
are  very  similar.  He  writes:  "Two  specimens  taken  '  in  cop.'  Settled 
on  under  side  of  a  twig,  heads  in  opposite  directions,  front  legs  of  each 
stretched  out  in  front  along  twig,  wings  laid  flat  along  abdomen." 

Styringomyia  didyma  Grimsh. 

1901    Styringomyia  didyma  Grimsh.     Fauna  Hawaiiensis,  p.  10. 

In  Fauna  Hawaiiensis,  Perkins  (19 13:  clxxxii)  mentions  the  breeding 
of  Styringomyia  didyma  by  F.  W.  Terry.  Before  the  reference  could  be 
investigated  by  the  writer,  Mr.  Terry  died.  The  following  letters  from 
O.  H.  Swezey  in  regard  to  the  matter  were  then  received. 

In  response  to  a  letter  sent  on  February  21,  1915,  Mr.  Swezey  replied 
on  March  26  that  "  no  information  on  the  rearing  of  the  species  is  available 
in  Mr.  Terry's  notes." 


THE  CRANE-FLIES  OF  NEW  YORK  — PART  II  959 

In  a  letter  dated  April  10  of  the  same  year,  however,  the  following 
notes  were  enclosed: 

About  forty-five  eggs  deposited  in  tube  December  5,  1910.  Chorion  jet  black,  shining  and 
thick,  resisting  dryness,  0.3  mm.  by  0.15  mm.,  very  finely  parallel-striate. 

One  hatched  December  10.  The  batch  was  placed  with  rotten  apples  and  cow  manure, 
hatching  December  15.  Larva  long,  head  small,  mandibles  distinct  and  well  chitinized! 

An  adult  male  emerged  about  January  21,  1911. 

Mr.  Swezey,  thru  whose  kindness  the  above  notes  are  available,  adds: 
"You  see  from  the  notes  of  Terry's  that  he  did  not  breed  Stynngomyia 
didyma  in  its  natural  habitat.  That  is  yet  unknown,  I  guess." 

The  striking  feature  of  this  life  history  is  its  brevity,  the  entire  egg, 
larval,  and  pupal  stages  being  passed  in  about  a  month  and  a  half.  The 
writer  knows  of  no  other  crane-flies  in  which  this  is  equaled,  its  nearest 
approach  presumably  being  in  the  smaller  Eriopterini. 

SUBFAMILY*  Cylindrotominae 

The  subfamily  Cylindrotominae  constitutes  a  small,  isolated  group  of 
crane-flies,  with  twenty  described  species  arranged  in  seven  recent  genera. 
All  the  species  are  Holarctic  in  their  distribution  with  the  exception  of 
five  species  of  the  Oriental  genera  Stibadocera  EnderL,  Stibadocerella 
Brun.,  and  Agastomyia  de  Meij.  The  group  is  a  decadent  one,  having 
been  much  better  developed  in  the  early  and  middle  Tertiaries  than  at 
present  (page  764). 

The  adult  flies  are  sluggish  in  their  habits,  occurring  on  vegetation 
in  cool,  shaded  spots.  The  species  of  Cylindrotoma  are  brightly  colored, 
yellow  and  black,  but  the  other  forms  are  somber  in  appearance  and 
black  or  dark  in  color,  the  body  being  in  some  cases  highly  polished  or 
metallic.  The  immature  stages  of  the  Cylindrotominae  differ  from  those 
of  all  other  Tipulidae,  so  far  as  is  known  to  the  writer,  in  being  spent 
on  various  bryophytic  and  spermatophytic  plants,  on  the  leaves  of  which 
the  larvae  feed.  The  larvae  are  usually  bright  green  in  color  and  suggest 
a  caterpillar  in  their  general  form.  Most  of  them  simulate  their  host 
plants  to  an  astonishing  degree.  The  immature  stages  of  Cylindrotoma 
and  Liogma  are  terrestrial,  while  those  of  Triogma  and  Phalacrocera  are 
aquatic  or  nearly  so. 

The  genera  of  the  subfamily  Cylindrotominae  may  be  separated  as 
follows : 


960  CHARLES  PAUL  ALEXANDER 

Larvae 

1.  Body  appendages  very  long  and  filiform;  aquatic Phalacrocera  Schin.  (p.  961) 

Body  appendages  short,  leaflike  or  tuberculate 2 

2.  Dorsal  appendages  all  simple,  on  the  terminal  abdominal  segments  in  a  single  row;  ter- 

restrial on  spermatophy tic  plants Cylindrotoma  Macq.  (p.  966) 

Dorsal  appendages  with  teeth  on  anterior  convex  side 3 

3.  Some  of  the  dorsal  appendages  with  three   or  four  teeth  on  anterior  face;   aquatic  on 

mosses Triogma  Schin.  (p.  973) 

Dorsal  appendages  with  cne  (L.  nodicornis)  or  two  (L.  glabrata}  teeth. 

Liogma  O.  S.  (p.  969) 

Pupae 

1.  Basal  abdominal  tergites  without  spines 2 

Basal  abdominal  tergites  with  acute  spines 3 

2.  Mesonotum  unarmed;  segments  6  and  8  each  with  two  powerful  dorsal  hooks;  segment 

7  with  a  pair  of  strong  ventral  spines;  pronotal  breathing  horns  elongate,  directed 

backward Phalacrocera  Schin.  (p.  961) 

Mesonotum  with  two  flattened  erect  lobe 3;  segments  6,  7,  and  8  naked;  pronotal  breathing 
horns  small,  directed  slightly  forward Cylindrotoma  Macq.  (p.  966) 

3.  Abdominal  spines  branched .Liogma  (glabrata)  (p.  669) 

Triogma  (trisulcata)  (p.  974) 
Abdominal  spines  not  branched Liogma  (nodicornis)  (p.  971) 

The  most  important  literature  on  the  Cylindrotominae  is  as  follows: 

General  account  of  subfamily.  Osten  Sacken,  1897;  Alexander,  1914:105-106;  Malloch> 
1915-17  b:  210-211;  Lenz,  1920  b:  113-115. 

Phalacrocera  replicala Larva,  pupa,  general. . .  De  Geer,  1773;  1776: 135-141,[351. 

Phalacrocera  replicata Larva Grube,  1868. 

Phalacrocera  replicata Larva Engel,  1884. 

Phalacrocera  replicata General Giard,  1895  b. 

Phalacrocera  replicata Larva,  general Bengcsson,  1897.  (Morphology  of 

larva.) 

Phalacrocera  replicata Larva,  pupa,  general  . .  Miall  and  Shelf ord,  1897.  (Mor- 
phology of  larva  and  pupa.) 

Phalacrocera  replicata Larva Bengtsson,  1899.  (Morphology  of 

heart.) 

Phalacrocera  replicata Larva Holmgren,  1908.  (Morphology  of 

head.) 

Phalacrocera  replicata Larva,  pupa Grunberg,  1910:32-35. 

Phalacrocera  replicata Larva,  pupa,  general. . .   Wesenberg-Lund,  1915:343-347. 

Phalacrocera  replicata Larva,  pupa,  general.  .  .   Lenz,  1920 b:  127-129. 

Cy'.indro'.oma  distinctissima. . . .   Larva,  general Schellenberg,  1803:22-23. 

Cylindrotoma  distinctissima..  .  .   Larva,  pupa,  general. ..   Boie,  1838:234. 

Cylindrotoma  distinctissima ....   Larva,  pupa,  general . . .   Zeller,  1842. 

Cylindrotoma  distinctissima. . .  .   General Schiner,  1864:563. 

Cylindrotoma  distinctissima. . .  .   Larva,  pupa,  general. . .   Kaltenbach,  1874:7. 

Cylindrotoma  distinctissima. . .  .  Larva Wesenberg-Lund,  1915:335  (as  Tri- 
ogma). 

Cylindrotoma  distinctissima.. .  .   Larva,  pupa,  general.  . .  Lenz,  1920  b:  115-117. 

Cylindrotoma  splendens Larva,  pupa,  general. . .   Cameron,  1918. 

Liogma  glabrata Larva,  general De  Rossi,  1876. 

Liogma  glabrata General Osten  Sacken,  1878  a. 


THE  JCRANE-FLIES  OF  NEW  YORK  —  PART  II  961 

Liogma  glabrata Larva,  pupa,  general . . .   Miiggenberg,  1901. 

Liogma  glabrata Larva,  general Da  Rossi,  1902. 

Liogma  glabrata General Alexander,  1914 : 106-107. 

Liogma  glabrata Larva,  pupa,  general.  . .   Lenz,  1920 b:  117-121. 

Liogma  nodicornis Larva,  pupa,  general. . .   Alexander,  1914: 107-115. 

Triogma  trisulcata Larva,  general Steinmann,  1907-08. 

Triogma  trisulcata Larva,  pupa,  general . .   Muller,  1908-09. 

Triogma  trisulcata Larva,  general Wesenberg-Lund,  1915:347-348  (as 

Liogma  glabrata). 

Triogma  trisulcata Larva,  pupa,  general..  .  Lenz,  1920 b:  121-127. 

Genus  Phalacrocera  Schiner  (Gr.  bald  +  horn) 

1863     Phalacrocera  Schin.     Wien.  Ent.  Monatschr.,  vol.  7,  p.  224. 

Larva.—  Body  covered  with  numerous  elongate,  trachea-bearing  filaments,  the  posterior 
pair  on  dorsal  segments  deeply  forked,  the  others  simple.  Spiracular  disk  with  dorsal  pair 
of  lobes  formed  by  rudimentary  posterior  branch  of  branched  filaments  of  eighth  abdominal 
segment.  Head  capsule  compact.  Mentum  with  about  fifteen  teeth. 

Pupa.' —  Cephalic  crest  low,  not  setiferous.  Pronotal  breathing  horns  long,  almost 
straight.  Dorsal  abdominal  segments  with  tubercles,  those  of  sixth  and  eighth  segments 
enlarged  into  spinous  hooks;  two  pointed  tubercles  on  seventh  sternite. 

Phalacrocera  is  a  small  genus  (four  species)  of  medium-sized  to  large, 
dull-colored  flies,  of  which  the  genotype,  Phalacrocera  replicata,  is  Euro- 
pean, P.  mikado  Alex,  is  Japanese,  and  the  two  remaining  species  are 
North  American. 

The  adult  flies  of  the  American  species  are  not  common,  the  best- known, 
P.  tipulina  0.  S.,  being  most  frequently  found  in  or  near  sphagnum  bogs  in 
mountainous  localities.  Needham  (1908  a :  209)  found  the  wings  of  an  indi- 
vidual of  this  species  in  the  pitcher  plant,  SarraceniapurpureaLmn.,  in  the 
Adirondack  Mountains,  together  with  the  wings  of  four  specimens  of  Ele- 
phantomyia  westwoodi  and  numerous  other  insects.  Most  of  the  specimens 
that  have  been  found  by  the  writer  were  taken  in  close  proximity  to  bogs. 

The  immature  stages  of  Phalacrocera  replicata  have  long  been  known, 
having  been  described  by  De  Geer  and  other  early  workers  on  insect 
biology.  More  recently  the  life  history,  anatomy,  and  morphology  have 
been  discussed  in  commendable  detail  by  several  other  writers  (page  960). 
Both  the  larvae  and  the  pupae  are  aquatic,  living  among  submerged 
plants  in  quiet,  but  non-stagnant,  water. 

The  immature  stages  of  P.  tipulina  are  very  much  to  be  desired,  as 
the  adult  shows  some  features  in  its  organization  not  found  in  the  other 
species  of  the  genus.6 

6  The  larva  of  this  species  was  discovered  by  J.  Speed  Rogers  in  1920. 


962  CHAKLES  PAUL  ALEXANDER 

Bengtsson  (1897)  erected  for  this  genus  the  group  Erucaeformia,  which 
he  considered  as  the  primitive  form  from  which  the  Nematocera  and  the 
Brachycera  have  been  derived.  This  group,  of  course,  has  no  standing 
whatsoever. 

Phalacrocera  replicata  (Linn.) 

1761     Tipula  replicata  Linn.    Fauna  Suecica,  2d  ed.,  p.  500-502. 

1863     Phalacrocera  replicata  Schin.     Wien.  Ent.  Monatschr.,  vol.  7,  p.  224. 

The  larvae  and  the  pupae  of  Phalacrocera  replicata  have  been  discussed 
in  such  detail  by  Miall  and  Shelf  ord,  by  Bengtsson,  by  Holmgren,  and 
by  Wesenberg-Lund,  that  they  are  considered  here  only  in  general  terms. 
The  habits  of  the  immature  stages  have  been  discussed  by  many  writers 
since  the  time  of  De  Geer.  They  are  oftentimes  rather  numerous  among 
aquatic  plants  such  as  Ranunculus  fluitans  Lam.,  Fontinalis  antipyretica 
Linn.,  Hypnum  elodes  Schp.,  H.  exannulatum  Guenbel,  and  other  species, 
feeding  on  these  mosses  and  probably  on  other  plants.  These  moss  frag- 
ments give  a  green  tinge  to  young  larvae  when  seen  thru  the  nearly  trans- 
parent body  wall.  Older  larvae  are  more  opaque  and  are  brownish  green 
in  color,  indistinctly  striped  with  pale  and  darker.  The  larva  is  extremely 
sluggish,  remaining  almost  motionless  for  hours.  It  clings  to  moss  stems 
by  its  large  anal  hooks,  and,  thus  secured,  it  often  sways  its  body  from 
side  to  side  as  if  to  accelerate  respiration.  The  larvae  can  go  for  long 
periods  of  time  without  fresh  air.  Miall  and  Shelford  kept  specimens 
alive  for  two  weeks  in  bottles  completely  filled  with  water,  and  for  a 
long  time  in  water  that  had  been  boiled.  The  larvae  can  live  for  a  long 
time  out  of  water.  Progression  thru  the  mats  of  submerged  vegetation 
is  accomplished  by  grasping  with  the  mandibles  and  the  anal  hooks, 
alternately.  When  alarmed  the  larvae  curl  into  a  rounded  ball,  after 
the  manner  of  many  caterpillars.  The  skin,  and  more  especially  the  long 
body  processes,  are  often  covered  with  ectoparasitic  organisms,  such 
as  algae  and  infusoria,  on  which  small  fresh-water  mollusks,  Plan- 
orbis,  have  been  observed  feeding  and  creeping  about  over  the  body  of 
the  larva.  This  coating  of  organisms,  the  body  outgrowths,  and  the  general 
coloration  of  the  larva,  give  it  a  striking  resemblance  to  the  mosses  among 
which  it  lives.  De  Geer  (1776:355)  shows  that  the  larva  can  endure 
excessive  cold.  He  placed  four  larvae  in  a  vessel  at  the  beginning  of 
winter,  and  examined  them  in  the  following  May.  During  the  winter 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  963 

the  water  in  which  the  larvae  lived  had  frozen  into  a  solid  mass,  yet, 
on  investigating  the  jar  in  the  spring,  De  Geer  found  two  of  the  larvae 
still  alive  and  able  to  feed,  and  within  a  month  both  had  pupated. 

The  larval  habitat  is  in  ponds  in  which  a  moss  vegetation  flourishes 
and  in  which  currents  keep  the  water  in  constant  motion.  A  female 
fly  was  observed  by  Miall  and  Shelford  (1897:360)  depositing  her  eggs 
in  the  leaf  axils  of  a  submerged  moss.  The  eggs,  about  sixty  in  number, 
are  laid  singly  and  adhere  slightly  to  the  moss;  they  are  opaque,  dark 
in  color,  and  spindle-shaped,  with  the  surface  of  the  chorion  irregularly 
pitted,  and  with  a  rosette-like  micropyle  at  one  end.  Bengtsson,  Mtiggen- 
berg,  and  others  believe  that  Phalacrocera  has  but  a  single  brood  in  a 
year,  the  larval  existence  occupying  about  eleven  months;  Miall  and 
Shelford,  however,  admit  the  possibility  of  a  second  brood.  The  egg 
stage  requires  from  eight  to  twelve  days  and  the  pupal  duration  is  seven 
or  eight  days,  according  to  Bengtsson.  According  to  Miall  and  Shelford, 
the  pupal  period  is  considerably  longer. 

During  the  larval  development  there  are  numerous  mol tings,  at  least 
eight  and  possibly  ten;  the  old  larval  skin  is  cast  by  a  simple  dorsal  split 
extending  from  the  first  to  the  fourth  segment.  Just  after  emerging 
from  the  egg  the  larva  is  from  2  to  2.25  millimeters  in  length  and  about 
0.5  millimeter  in  diameter,  excluding  the  body  projections.  The  first 
larval  stage  (Bengtsson,  1897)  lasts  until  the  second  molting.  The  body 
is  provided  with  ten  pairs  of  long,  delicate,  threadlike,  lateral  projections, 
which  are  located  on  the  second  to  the  eleventh  body  segments  and  are 
half  as  long  as  the  body.  The  other  projections  of  the  older  larvae  are 
merely  indicated.  The  attachment  apparatus  is  placed  immediately 
before  the  anus,  and  consists  of  from  eight  to  twelve  chitinized  hooks, 
directed  forward  and  arranged  in  an  arcuated  crossrow.  The  color  of 
the  body  is  white,  almost  transparent.  The  mouth  parts  show  the  mandi- 
bles without  a  prostheca  and  moving  horizontally.  The  second  larval 
stage  lasts  from  the  second  until  the  fourth  molting.  The  body  appendages 
have  appeared  and  are  clearly  developed,  resembling  in  appearance  and 
relative  length  those  of  the  definitive  stage.  The  attachment  apparatus 
is  post-anal.  The  body  takes  on  a  distinctly  striped  appearance.  The 
mouth  parts  have  the  prostheca  well  developed  on  the  mandibles,  which 
are  vertically  placed  and  therefore  have  an  up-and-down  movement. 
The  third  larval  stage  represents  the  fuUy  grown  larva  from  the  fourth 


964  CHARLES  PAUL  ALEXANDER 

molt  up  to  the  time  of  pupation.  Here  the  dorsum  of  the  body  is  a  dirty 
brownish  green  with  more  or  less  distinct  brighter  spots,  and  the  venter 
is  bright  green  in  color.  The  mouth  parts  and  the  attachment  apparatus 
are  as  in  the  second  stage. 

The  pupa  is  comparatively  active,  moving  about  by  flexion  of  the 
abdomen.  Its  usual  position  is  vertical,  with  the  tips  of  the  breathing 
horns  just  reaching  the  surface  of  the  water.  This  vertical  position  the 
pupa  maintains  by  grasping  the  vegetation  with  its  caudal  abdominal  hooks. 
At  times  the  pupa  descends  beneath  the  water  by  clinging  to  the  vegetation, 
but  a  submergence  of  six  hours  causes  asphyxiation.  The  pupal  existence 
was  determined  by  Miall  and  Shelford  as  eleven  days.  When  the  adult 
emerges,  the  cast  pupal  skin  is  left  attached  to  a  moss  leaf  by  the  dorsal 
abdominal  projections  at  the  posterior  end  of  the  body. 

Larva. —  Length,  about  25  mm. 

Young  larvae  distinctly  greenish,  especially  on  ventral  side,  this  coloration  caused,  at 
least  in  part,  by  contents  of  alimentary  canal  showing  thru  body  wall;  older  larvae  more 
opaque,  brownish  green  in  color;  dorsum  with  an  indistinct  striping  of  brown  and  whitish; 
ventral  surface  whitish. 

Head  entirely  retractile  within  prothorax  and  usually  so  retracted  except  when  larva  is 
feeding;  opening  transverse.  Prothorax,  viewed  from  above,  roughly  rounded,  anterior 
margin  convex;  on  ventral  surface  traversed  by  a  weak  suture.  Meso-  and  metathorax 
narrow.  Abdominal  segment  1  indistinctly  divided  into  two  annuli,  the  more  basal  one 
very  narrow;  abdominal  segments  2  to  7  divided  into  a  narrow  basal  ring  and  a  much  broader 
posterior  ring,  each  of  these  annuli  still  further  subdivided  into  two  annulets.  Body  pro- 
vided with  numerous  elongate  trachea-bearing  filaments,  both  simple  and  branched,  giving 
larva  a  very  bristly  appearance  (Plate  LXXX.III,  448) ,  these  spines  arranged  as  follows :  tergites 
with  both  simple  and  bifurcated  filaments;  on  posterior  part  of  pronotum  two  short,  simple 
filaments;  on  meso-  and  metanotum,  two  pairs  of  simple  filaments;  on  abdominal  segment 
1,  an  anterior  pair  of  simple,  and  a  posterior  pair  of  deeply  branched,  filaments;  segments 
2  to  7  with  basal  ring  unarmed,  posterior  ring  with  an  anterior  pair  of  simple,  and  a  caudal 
pair  of  deeply  branched,  filaments;  .segment  8  with  only  a  branched  pair,  anterior  branch 
long,  slender,  posterior  branch  very  small,  its  outer  face  heavily  chitinized  and  forming 
dorsal  lobes  of  spiracular  disk;  pleurites  with  all  the  filaments  simple;  one  on  posterior  part 
of  prothorax,  and  on  anterior  part  of  each  of  the  other  two  thoracic  segments;  posterior 
filament  on  these  latter  nearly  vestigial;  two  unequal  filaments  on  first  abdominal  segment; 
segments  2  to  7  with  one  filament  on  basal  ring  and  two  on  posterior  ring,  the  anterior  one 
the  longest;  segment  8  with  a  single  rudimentary  filament;  sternites  with  all  the  filaments 
simple;  prosternum  without  filaments;  meso-  and  metasternum  with  a  strong  filament  near 
lateral  margins;  abdominal  segment  1  with  two  pairs  of  filaments,  posterior  pair  the  longer 
and  more  widely  separated;  segment  2  with  three  pairs  of  filaments,  anterior  pair  very  short; 
segments  3  to  7  with  four  pairs  of  filaments  and  an  additional  median  one,  arranged  as  follows: 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  965 

two  pairs  of  small  filaments  on  basal  ring,  the  anterior  pair  more  widely  separated,  two 
pairs  of  much  longer  filaments  on  posterior  ring,  the  last  pair  more  widely  separated,  longer, 
and  tipped  with  blackish,  between  them  a  tiny  median  filament.  Chaetotaxy  as  follows: 
tergites  with  four  solitary  setae  along  anterior  margin  of  prothorax,  and  two  setae  just 
laterad  of  each  dorsal  filament  of  prothorax  and  posterior  filaments  of  meso-  and  meta thorax; 
abdominal  segments  with  a  seta  laterad  of  each  simple  and  branched  filament;  pleurites 
with  a  stiff  seta  at  base,  and  out  toward  apices,  of  middle  lateral  filaments;  sternites  with  a 
group  of  about  four  stiff  setae  on  either  side  of  posterior  ring  of  prosternum,  and  a  single 
stiff  seta  on  either  side  of  median  line;  meso-  and  metasternum  with  a  lateral  group  of  setae; 
a  seta  laterad,  and  another  at  about  midlength,  of  each  of  the  posterior  pair  of  ventral 
filaments. 

Spiracular  disk  with  ventral  lobes  elongate,  slightly  recurved,  the  posterior  face  intensely 
blackened,  chitinized,  each  lobe  with  two  acute  spines  at  tip,  with  two  setae  near  them; 
a  stiff  seta  on  side  of  base  of  each  ventral  lobe;  dorsal  lobes  as  already  described,  the  rudi- 
mentary posterior  branch  of  last  furcate  dorsal  filament  much  smaller  than  in  ventral  lobes, 
the  posterior  face  heavily  blackened;  spiracular  disk  rhombic  in  form,  white,  and  having 
almost  the  appearance  of  porcelain.  Spiracles  situated  between  bases  of  dorsal  lobes.  Skin 
about  spiracles  capable  of  retraction  so  as  to  form  .a  deep  recess. 

Head  capsule  rather  short,  almost  conical,  formed  of  two  large  lateral  plates  and  a  some- 
what smaller  and  shorter  prefrontal  plate.  Mentum  with  an  outer  (ectolabial)  part  and 
an  inner  (endolabial)  part;  mentum  a  triangular  or  somewhat  pentagonal  plate,  strongly 
chitinized,  fore  margin  with  about  fifteen  teeth;  mandibles  working  against  teeth  of  both 
endo-  and  ectolabia.  Antenna  of  a  single  segment,  bearing  on  its  truncated  apical  end  a 
few  sensory  papillae.  Mandibles  small  but  strong,  curved  inward  at  tip  and  furnished  with 
a  fringe  of  setae,  which  assist  in  closing  the  mouth  opening.  Maxilla  expanded  into  flattened, 
shovel-like  structures,  inserted  high  on  side  of  head;  palpus  with  a  number  of  sensory  papillae 
at  apex,  and  with  a  porous  plate  on  outer  side  which  seems  to  be  an  organ  of  hearing. 

Pupa. —  Length,  16-18  mm.     (Miall  gives  length  up  to  20  mm.) 
Width,  d.-s.,  2.9-3.3  mm. 
Depth,  d.-v.,  2.7-2.9  mm. 

Coloration  greenish  brown,  in  alcohol  a  paler  yellowish  brown;  a  very  broad  dorso-median 
dark  brown  stripe  which  is  narrowly  margined  laterally  with  yellowish;  dorsum  of  abdomen 
suffused  sublaterally  with  dusky,  extreme  lateral  margins  of  body  yellowish;  ventral  surface 
with  two  broader  sublateral  stripes  and  a  very  narrow  ventro-median  stripe. 

Anterior  end  of  body  very  deep  and  thick,  as  is  usual  in  this  group  of  crane-flies.  Abdomen 
greatly  depressed,  with  lateral  margins  very  thin  and  flattened.  Cephalic  crest  low,  non- 
setiferous,  located  between  antennal  bases.  Labrum  broad  basally,  narrowed  toward  apex, 
which  is  broadly  rounded;  two  setae  at  base  of  labrum.  Labial  lobes  subcircular  in  outline. 
Maxillary  palpi  elongate,  bent  strongly  backward  so  as  to  lie  along  flattened  cheek.  Antenna 
rather  elongated,  extending  far  beyond  origin  of  wing  pad  (Plate  LXXXIII,  449).  Thorax 
very  deep,  flattened  above.  Pronotal  breathing  horns,  which  are  broken  in  the  writer's  speci- 
mens, seen  from  other  descriptions  to  be  rather  elongate,  almost  straight,  and  slightly 
divergent;  two  small  setae  between  bases  of  breathing  horns;  two  groups  of  setae  on  anterior 
part  of  pronotum  before  breathing  horns.  Mesonotum  (Plate  LXXXIII,  449)  with 


six 


966  CHARLES  PAUL  ALEXANDER 

small,  setiferous  tubercles,  one  above  each  wing  axil,  the  other  four  arranged  in  a  trapezoid 
on  dorsum  with  the  anterior  pair  closer  together;  metanotum  with  six  setiferous  tubercles, 
four  of  which  are  median  in  position,  the  other  pair  at  antero-lateral  margin,  near  base  of 
halteres.  Wing  sheaths  ending  about  opposite  apex  of  second  abdominal  segment.  Leg 
sheaths  ending  before  apex  of  third  abdominal  segment. 

First  abdominal  segment  similar  to  metanotum;  segments  2  to  7  indistinctly  subdivided 
into  a  narrow  basal  ring  and  a  much  broader  posterior  ring;  basal  ring  unarmed  except  for 
a  seta  on  pleural  margin  and  a  trapezoid  of  tubercles  on  sternum;  posterior  annulus  armed 
as  follows:  tergites  with  four  tubercles  arranged  in  a  quadrangle  and  located  in  the  dark 
median  stripe,  posterior  tubercles  with  a  stiff  seta  just  laterad  of  each;  a  lateral  seta  on  a 
line  with  anterior  tubercles;  two  lateral  setae  on  a  level  with  posterior  tubercles,  the  proximal 
one  considerably  the  larger;  these  setae  located  in  the  yellow  sublateral  stripe;  pleural  margin 
with  two  setiferous  tubercles  with  an  acute  subappressed  spine  between  them;  a  seta  immedi- 
ately ventrad  of  anterior  tubercle;  on  segments  6  and  7,  spine  closer  to  caudal  margin  of 
segment,  and  tubercle  beyond  it  lacking;  sternites  with  a  trapezoid  of  naked  tubercles  on 
basal  ring,  posterior  pair  the  closer  together;  a  trapezoid  of  larger  setiferous  tubercles  on 
posterior  ring,  anterior  pair  the  closer  together;  posterior  punctures  each  having  two  setae, 
with  an  additional  slender  seta  laterad  of  these  in  the  lateral  dark  stripe;  dorsum  of  segment 
6  (Plate  LXXXIII,  450)  with  posterior  pair  of  tubercles  replaced  by  two  powerful  lobes 
directed  caudad  and  laterad;  segment  7  unarmed  on  dorsum,  sternum  with  two  acute  spines 
near  caudal  margin;  segment  8  with  lateral  angles  produced  dorsad  into  slender  lobes  which- 
are  spinous  on  all  the  faces;  caudal  angles  directed  caudad  into  slender  lobes,  acute  at  tips 
and  with  anterior  inner  face  spinous  and  bearing  a  few  setae.  Male  cauda  with  sternal 
valves  very  blunt  and  rounded,  on  either  side  of  ventro-median  line  produced  caudad  into  an 
acute  spine.  Female  cauda  with  dorsal  acidothecae  a  little  longer  than  the  more  slender 
sternal  valves,  both  pairs  much  exceeded  by  caudal  angles  of  eighth  segment. 

Nepionotype. —  Larva,  Denmark. 

Neanotype. —  Pupa,  Denmark. 

Paratypes. —  One  larva  and  one  pupa. 

(The  writer  is  indebted  to  Dr.  C.  Wesenberg-Lund  for  this  material, 
as  well  as  for  several  other  interesting  European  crane-fly  life  histories.) 

Genus  Cylindrotoma  Macquart  (Gr.  cylinder  4-  /  cut) 

1834    Cylindrotoma  Macq.    Suit,  a  Buff.,  vol.  1,  Hist.  Nat.  Ins.,  Dipt.,  p.  107. 

Larva. —  Body  covered  with  simple  tubercles,  a  median  dorsal  row  and  a  double  ventral 
row.  Spiracular  disk  large,  surrounded  by  small  lobes.  Head  capsule  compact.  Mentum 
with  about  fifteen  teeth. 

Pupa.' —  Pronotal  breathing  horns  short,  cylindrical,  directed  strongly  ventrad.  Meso- 
notum  with  two  flattened  lobes  directed  cephalad.  Abdomen  unarmed  with  spines  or  lobes. 

Cylindrotoma  is  a  small  genus  including  six  species  distributed  thruout 
the  North  Temperate  Zone,  three  of  these  species  occurring  in  North 
America. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  967 

The  adult  flies  of  Cylindrotoma  tarsalis  Johns.,,  the  only  species  that 
the  writer  has  ever  found  in  nature,  are  common  on  rank  vegetation  in 
cool  boggy,  and  swampy  woods.  The  immature  stages  of  the  genotype 
C.  d^st^nct^ss^ma  Meig ;.),  have  long  been  known,  having  been  discussed 
by  Schellenberg  (1803),  Boie  (1838),  Zeller  (1842),  Kaltenbach  (1874), 
and  others.  The  larva  shows  some  resemblance  to  that  of  Phalacrocera 
but  is  almost  entirely  terrestrial  in  its  habits,  feeding  on  the  leaves  of 
various  spermatophytic  plants  such  as  Caltha  palustris,  Anemone  nemorosa 
Ranunculus  repens,  Chrysosplenium,  Stellaria  nemorum,  Sanicula  europaea 
Viola  biflora,  Valeriana  officinalis,  Allium,  and  others.  It  attains  a  length 
of  nearly  25  millimeters,  and  is  narrow,  depressed,  tapering  to  either  end 
and  of  a  grass-green  color.  There  is  a  slight  dorsal  ridge  from  which  a 
row  of  short,  fleshy  spines  projects,  these  spines  being  directed  backward 
and  one  spine  on'  each  segment  being  longer  than  the  others.  There  is  a 
broad  lateral  margin  bearing  very  short  processes,  and  there  are  also 
eight  pairs  of  ventral  ridges  without  hooks  and  a  pair  of  longer  backward- 
directed  processes  near  the  anus.  The  pupa  affixes  itself  to  stalks  or  leaves 
by  the  caudal  end,  to  which  the  remains  of  the  last  larval  skin  adhere. 
The  larvae  generally  remain  on  the  lower  surface  of  the  leaves,  on  which 
they  feed,  gnawing  holes  in  them.  When  about  to  pupate  they  generally 
leave  their  food  plants  and  fasten  themselves  on  grass  blades  and  leaves 
near  by,  usually  pupating  the  following  day.  From  the  foregoing  obser- 
vations it  would  seem  that  there  are  in  the  various  localities  two  genera- 
tions a  year,  one  in  the  spring  and  the  other  in  the  autumn. 

The  life  history  of  C.  spkndens  has  recently  been  worked  out  in  con- 
siderable detail  by  Dr.  A.  E.  Cameron,  thru  whose  kindness  the  writer 
has  received  specimens  for  study. 

Cylindrotoma  spkndens  Doane 

1900     Cylindrotoma  spkndens  Doane.     Joura.  N.  Y.  Ent.  Soc.,  vol.  8,  p.  197. 

Cylindrotoma  juncta  Coq.     Proc.  Wash.  Acad.  Sci.,  vol.  2,  p.  401. 
1918     Cylindrotoma  spkndens  Cameron.    Ann.  Ent.  Soc.  Amer.,  vol.  11,  p.  67-89. 

Cylindrotoma  spkndens  is  an  interesting  crane-fly  occurring  from  British 
Columbia  northward  to  Alaska.  Dr.  Cameron's  excellent  notes  on  the 
life  history  of  this  species  are  abstracted  in  detail  on  pages  708  to  710 
of  this  paper.  The  following  descriptions  were  made  from  material  sent 


968  CHARLES  PAUL  ALEXANDER 

to  the  writer  by  Dr.  Cameron,  a  few  details  being  added  from  Dr.  Cameron's 
published  notes. 

Larva. —  Length,  15-17  mm. 

Width,  d.-s.,  2.4-2.5  mm. 
Depth,  d.-v.,  1.5  mm. 

Coloration  light  chlorophyll  green,  with  two  narrow,  pale  brown  lines  on  dorsum,  extending 
from  posterior  end,  above  spiracles,  anteriorly,  becoming  more  expanded  and  diffused  on 
fore  part  of  body. 

Body  very  depressed,  both  dorsal  and  ventral  surfaces  being  flattened,  lateral  margins 
sharp  (Plate  LXXXIV,  453).  Head  completely  retractile  within  prothorax.  Skin  delicately 
reticulated  and  roughened.  Thorax,  viewed  from  above,  semicircular  in  outline,  margined 
with  about  four  tubercles  on  either  side,  anterior  pair  the  largest,  separated  by  a  V-shaped 
notch.  Pronotum  with  an  anterior  pair  of  small  tubercles  and  a  larger  median  tubercle 
behind,  directed  backward;  segments  2  and  3  each  with  lateral  margins  two- toothed,  the 
anterior  tooth  the  larger  and  more  pointed;  a  blunt  dorsal  tubercle  on  anterior  margin  and 
a  larger  one  behind.  Abdominal  segments  indistinctly  divided  into  four  annuli  which  are 
poorly  delimited;  the  two  basal  annuli  narrow  and  corresponding  to  basal  annulus  of  most 
crane-fly  larvae,  the  third  annulus  the'  largest  and  bearing  a  slender  lateral  tooth;  other 
segments  less  distinctly  divided  serrately  on  lateral  margin;  dorsum  of  abdominal  segments 
with  a  row  of  blunt  median  tubercles,  there  being  four  on  each  segment,  the  first  very  small 
to  vestigial,  the  third  slender,  the  last  the  stoutest;  on  .venter  of  abdominal  segments  1  to 
7,  near  posterior  margin,  a  fleshy  conical  lobe  (pseudopodium)  on  either  side  of  median  line, 
these  being  smaller  on  anterior  segments  (1  to  4),  and  much  larger  and  paler  on  posterior 
segments  (5  to  7).  Spiracular  disk  very  large  and  flattened,  obliquely  truncated,  surrounded 
by  six  small  lobes,  the  dorso-lateral  pair  small,  widely  separated,  the  ventro-lateral  pair 
much  larger,  the  ventral  pair  very  small,  slender.  Spiracles  very  small,  circular,  widely 
separated,  the  distance  between  them  about  equal  to  six  times  diameter  of  one. 

Mouth  parts  with  labrum  linguliform,  terminating  in  four  teeth,  external  pair  smaller 
than  internal  pair.  Mentum  with  seven  teeth  on  either  side  of  small  median  one,  the  first 
and  the  third  on  either  side  being  the  largest.  Hypopharynx  with  two  rows  of  small  teeth. 
Antenna  two-segmented;  basal  segment  elongate,  pyriform;  apical  segment  thimble-shaped. 
Mandible  (Plate  LXXXIV,  454)  powerful;  cutting  edge  with  about  five  marginal  teeth  and 
two  dorsal  teeth;  a  large  basal  prostheca.  Maxilla  large;  outer  lobe  fringed  with  stiff  hairs; 
a  few  acute  sensory  bristles  situated  on  elevated  papillae,  two  on  cardo,  one  on  outer  lobe 
of  maxilla;  maxillary  palpi  short,  cylindrical,  apex  obliquely  truncated  and  provided  with 
numerous  sensory  knobs,  on  side  near  tip  a  circular  porous  plate  (which  as  it  appears  in 
Phalacrocera  is  suggested  by  Bengtsson  as  probably  being  an  organ  of  hearing). 

Pupa. —  Length,  11.7-14  mm. 

Width,  d.-s.,  2-2.8  mm. 
Depth,  d.-v.,  1.4-2  mm. 

Color  of  live  pupa,  leaf  green;  preserved  specimens  much  paler,  yellowish;  pronotal 
breathing  horns  grayish  white;  margins  of  abdomen  nearly  translucent. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  969 

Labrum  narrow,  apex  evenly  rounded.  Labial  lobes  stout,  separated  by  labrum,  at  tip 
narrowed  and  somewhat  pointed.  Maxillary  palpi  long  and  stout,  just  beyond  base  bent 
strongly  backward.  Antennae  rather  short,  bases  not  widely  separated  (Plate  LXXXIV 
456).  Thorax  very  deep,  much  flattened  anteriorly  (Plate  LXXXIV,  455).  Pronotal 
breathing  horns  short,  cylindrical,  slightly  divergent,  apex  of  each  a  little  expanded.  Meso- 
notum  at  declivity  with  two  flattened  plates,  which  are  blunt  and  directed  cephalad  and 
slightly  laterad;  above  wing  axil  a  smaller  tubercle.  Wing  sheaths  ending  just  before  apex 
of  second  abdominal  segment.  Leg  sheaths  ending  just  before  apex  of  third  abdominal 
segment;  tarsi  ending  about  on  a  level. 

Abdomen  strongly  depressed,  lateral  margins  carinate;  segments  distinct,  unarmed,  sub- 
divided into  a  narrow  basal  ring  and  a  much  broader  posterior  ring;  segments  2  to  5  a  little 
longer  than  segments  6  and  7.  Male  cauda  with  valves  divided  into  two  rounded  lobes 
by  a  small  median  notch.  Female  cauda  with  dorsal  valves  straight,  a  little  longer  and 
much  stouter  than  the  slender  sternal  valves. 

Nepionotype. —  Westhclme,  Vancouver  Island,  B.  C.,  May  15,  1917. 
Neanotype. —  Female  pupa  with  type  larva. 

Genus  Liogma  Osten  Sacken  (Gr.  smooth  +  furrow) 
1869    Liogma  0.  S.    Mon.  Dipt,  N.  Amer.,  part  4,  p.  298. 

Larva.—  Body  covered  with  elongate,  leaflike  projections,  dorsal  ones  with  one  or  two 
teeth  on  anterior  convex  face.  Spiracular  disk  surrounded  by  four  lobes.  Head  capsule 
compact.  Mentum  with  about  fourteen  teeth. 

Pupa.' —  Metanotum  and  abdominal  tergites  with  elongate  spines,  branched  in  L.  glabrata, 
simple  in  L.  nodicornis. 

The  genus  Liogma  includes  three  known  species,  of  which  the  genotype, 
Liogma  nodicornis  (O.  S.),  is  American.  The  adult  flies  of  this  species 
are  sluggish,  and  are  found  resting  on  vegetation  growing  in  and  about 
shaded  ponds  and  similar  situations. 

In  Europe  the  life  history  of  L.  glabrata  (Meig.)  has  been  worked  out 
by  De  Rossi  (1876),  by  Wesenberg-Lund  (1915:347-348),  and  in  con- 
siderable detail  by  Miiggenberg  (1901).  The  larvae  were  found  in  the 
woods  near  Berlin,  in  wet,  grassy  spots  where  the  mossHypnum  squarrosum 
Brch.  &  Schp.  occurs.  The  complete  metamorphosis  of  the  insect  takes 
one  year,  the  larval  life  requiring  the  greater  part  of  this  period.  The 
egg  stage  lasts  from  eight  to  ten  days  and  the  pupal  stage  from  eleven  to 
twelve  days.  The  duration  of  adult  life  is  not  known,  but  it  is  certainly 
short,  occupying  but  a  few  weeks  at  the  most. 

Near  Berlin  the  flies  emerge  during  the  first  half  of  July.  The  males 
appear  first,  the  females  later,  and  the  latter  are  always  seized  in  copulation 


970  CHARLES  PAUL  ALEXANDER 

by  the  males  just  after  they  have  forsaken  the  pupal  skin  and  while  still 
teneral  and  undeveloped.  Each  female  lays  about  sixty  eggs,  and  these 
are  deposited  singly  on  the  leaves  or  branches,  or  attached  lightly  to  the 
axils  of  the  leaves,  of  Hypnum  squarrosum.  Egg  deposition  begins  from 
one-half  to  one  day  after  copulation,  and  may  be  extended,  with  many 
interruptions  of  greater  or  less  extent,  thru  a  whole  day.  The  eggs  are 
from  1  to  1.2  millimeters  long,  spindle-shaped,  yellowish  green  in  color, 
the  chorion  with  a  reticulate  sculpturing.  After  the  accomplishment  of 
oviposition  the  exhausted  female  soon  dies.  The  larvae  when  first  hatched 
are  2  millimeters  in  length  and  do  not  yet  possess  the  beautiful  moss-green 
color  of  the  later  stage.  With  the  exception  of  the  chitinized  head  capsule, 
they  appear  ashy  gray.  The  numerous  thornlike  projections  are  to  be 
noted  already  in  the  same  positions  as  those  of  the  developed  larva,  but 
do  not  show  the  branching  of  the  later  stages.  In  the  autumn  the  young 
larva  grows  very  slowly,  and  during  the  winter  it  is  still  very  small  and 
difficult  to  detect.  In  the  spring  the  growth  is  greatly  accelerated,  and 
the  larva  becomes  fully  grown  during  the  latter  half  of  June.  While 
attaining  its  growth  the  larva  molts  several  times,  probably  at  least 
eight  —  the  number  determined  for  Phalacrocera  by  Bengtsson.  Pupation 
occurs  in  the  moss  where  the  larva  happens  to  be.  In  its  green  color, 
with  brown  blotches,  the  larva  remarkably  simulates  its  host  plant  and 
the  effect  of  the  shadows  cast  by  the  plant  stems  and  leaves.  The  larvae 
are  extremely  sluggish  in  their  habits. 

The  American  species  Liogma  nodicornis  has  been  found  in  various  mosses 
of  the  genus  Hypnum  (Alexander,  1914).  The  immature  stages  of  this 
species  are  discussed  below. 

The  following  keys  separate  the  species  of  Liogma: 

Larvae 

Prothoracic  segment  bearing  four  conspicuous  dorsal  projections  about  on  a  line;  meso-  and 
metathorax  with  two  pairs  of  dorsal  appendages,  each  bearing  two  lateral  teeth  in  front; 
second  abdominal  segment  with  four  dorsal  appendages,  the  last  two  bearing  two  teeth 
in  front  (Palaearctic) glabrata  (Meig.)  (p.  969) 

Prothoracic  segment  bearing  four  inconspicuous  dorsal  tubercles;  meso-  and  metathorax  with 
two  pairs  of  dorsal  appendages,  the  anterior  pair  small,  both  pairs  simple;  second  abdominal 
segment  with  four  dorsal  appendages,  the  last  two  bearing  a  single  small  tooth  in  front 
(Nearctic) nodicwnis  (O.  S.)  (p.  971) 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  971 

Pupae 

Pronotal  breathing  horns  directed  cephalad  and  dorsad;  mesonotum  bearing  two  pairs  of 
spines,  the  more  anterior  being  the  smaller,  situated  just  behind  breathing  horns,  posterior 
pair  the  larger;  metanotum  with  two  pairs  of  spines;  abdomen  with  first  tergite  bearing 
two  pairs  of  spines,  the  first  having  two  lateral  branches,  the  second  simple;  second  tergite 
bearing  two  pairs  of  spines,  the  first  with  two  lateral  branches,  the  second  with  one  branch; 
third  tergite  bearing  three  pairs  of  spines,  the  first  very  short  and  simple,  the  second  with 
two  lateral  branches,  the  third  with  one  branch;  tergites  4  and  5  with  three  pairs  of  branches, 
the  first  two  similar  to  those  of  third  segment,  the  last  possessing  two  lateral 
branches glabrata  (Meig  )  (p.  969) 

Pronotal  breathing  horns  directed  cephalad  and  ventrad;  mesonotum  spineless;  metanotum 
with  one  pair  of  spines;  abdominal  tergites  bearing  but  a  single  pair  of  appendages,  which 
are  unbranched  and  correspond  in  position  to  the  last  or  more  posterior  of  those  of  the 
European  species nodicornis  (O.  S.)  (p.  971) 

Liogma  nodicornis  (0.  S.)  ^ 

1865     Triogma  nodicornis  0.  S.     Proc.  Ent.  Soc.  Phila.,  vol.  4,  p.  239. 
1869     Cylindrotoma  nodicornis  O.  S.     Mon.  Dipt.  N.  Amer.,  part  4,  p.  301. 
1887    Liogma  nodicornis  0.  S.    Berl.  Ent.  Ztschr.,  vol.  31,  p.  226. 

Liogma  nodicornis,  the  only  American  species  of  the  genus,  is  widely 
distributed  thruout  the  Northeastern  States.  The  larvae  are  found  in 
moss  —  Hypnum  cupressiforme  Linn,  and  related  species.  They  are  the 
most  sluggish  of  any  crane-flies  known  to  the  writer,  moving  only  with 
great  slowness  and  much  of  the  time  appearing  to  be  quite  dead.  They 
crawl  about  among  the  stems  of  their  host  plant  and  probably  never  leave 
it,  even  to  pupate.  The  pupal  duration  indoors  is  not  more  than  six 
days.  Further  details  of  the  life  history  are  given  by  the  writer  in  an 
earlier  paper  (Alexander,  1914). 

Larva. —  Length,  14.5-15  mm. 
Width,  3  mm. 
Depth,  2.5  mm. 

Color  of  live  larva  light  green,  the  numerous  spines  covering  the  body  darker;  sides  with 
seven  black  marks,  the  first  on  first  abdominal  segment,  the  last  on  seventh  abdominal 
segment;  the  marks  on  ends  the  smallest  and  least  distinct,  the  five  intermediate  marks 
large  and  conspicuous;  these  marks  all  lying  parallel  to  one  another;  posterior  face  of  ventral 
lobes  surrounding  stigmal  field  intensely  black. 

Prothorax  in  front  (Plate  LXXXV,  462)  sloping  from  anterior  end,  on  ventral  slope  provided 
with  liplike  lobes,  with  a  transverse  slit  from  which  head  capsule  is  exserted;  upper  lip  the 
higher,  not  strongly  chitinized,  provided  with  a  few  small,  scattered  bristles,  these  more 
numerous  on  sides  of  lobe;  lower  lip  not  so  high,  with  small,  scattered  bristles  not  arranged 
in  a  row  as  in  L.  glabrata:  at  angle  of  slit  a  small  rounded  lobe  bearing  a  small  bristle.  Dorsal 
body  appendages  reduced  to  a  pair  of  lobes  in  front,  separated  by  a  space  a  little  greater 
than  diameter  of  one,  and  a  pair  of  smaller  lobes  behind,  very  widely  separated;  lateral  body 
appendages  long,  conspicuous;  ventral  body  appendages  not  apparent.  Meso-  and  meta- 
thorax  swollen  and  arched  ventrally  like  prothorax;  dorsal  appendages  two,  a  small  conical 


972  CHARLES  PAUL  ALEXANDER 

one  in  front  and  a  much  larger  one  behind  bearing  a  small  tooth  in  front  and  with  its  tip 
directed  backward;  lateral  appendages,  viewed  from  above,  two  in  number,  anterior  one  the 
larger,  directed  sharply  backward,  the  second  smaller,  conical;  ventral  appendages,  viewed 
from  side,  four,  anterior  pair  the  larger,  posterior  pair  small,  slightly  behind  the  others. 

Dorsal  appendages  of  abdominal  segments  (Plate  LXXXV,  457)  as  follows:  first  segment  with 
two  pairs  of  appendages;  anterior  pair  the  shorter,  conical,  tips  strongly  recurved  and  bearing 
a  tiny  tooth  on  anterior  face  at  about  midlength;  posterior  pair  much  longer,  with  tips 
bent  strongly  backward,  a  small  tooth  on  anterior  face  at  about  one-third  length  from  base; 
segments  2  to  7  with  four  pairs  of  appendages,  the  first  pair  very  small,  conical,  the  second 
exactly  similar  but  larger,  the  third  and  fourth  pairs  similar  to  appendages  of  first  abdominal 
segment;  the  tiny  anterior  appendage  largest  on  second  segment,  gradually  becoming  smaller 
toward  end  of  body.  Lateral  abdominal  appendages  as  follows:  first  segment  with  three 
pairs  of  appendages,  the  first  directed  laterad,  the  posterior  two  more  recurved  and  directed 
caudad;  segments  2  to  7  with  four  pairs  of  appendages,  the  first  very  small,  situated  at  antero- 
lateral  angle  of  segment,  the  other  three  subequal  and  directed  caudad.  Ventral  abdominal 
appendages  as  follows:  first  segment  with  three  pairs  of  appendages,  which  are  successively 
larger  from  the  short  anterior  one  to  the  large  posterior  one;  segments  2  to  7  with  five  pairs 
of  appendages,  the  first  three  small,  the  fourth  intermediate  in  size  between  them  and  the 
enlarged  fifth  pair.  Eighth  segment  bearing  spiracular  disk  and  its  lobes;  dorsal  side  of 
field  with  a  pair  of  long,  slender  lobes  bent  conspicuously  cephalad.  Spiracular  disk  (Plate 
LXXXV,  461)  small,  oval,  the  two  rounded-oval  spiracles  situated  side  by  side  and  close  to  each 
other,  inclined  toward  each  other  and  capable  of  being  closely  appressed;  ventral  lobes  of 
disk  directed  ventrad,  inner  faces  of  lobes  with  a  conspicuous  jet-black  line,  tip  of  each  lobe 
ending  in  a  sharp  recurved  hook.  (Miiggenberg  regards  these  lobes  as  representing  the 
ninth  abdominal  segment.)  Ventral  surface  of  terminal  segments  with  small  protuberances. 

Head  completely  retractile  into  first  thoracic  segment.  Mentum  (Plate  LXXXV,  458)  with 
about  seven  teeth  on  either  side,  terminal  pair  the  larger,  separated  by  a  V-shaped  notch. 
Antenna  (Plate  LXXXV,  460)  twro-segmented,  basal  segment  elongate-cylindrical,  tip  very 
short,  thimble-shaped,  with  a  diameter  less  than  that  of  elongate  basal  segment.  Mandible 
(Plate  LXXXV,  459  and  460)  many-toothed  on  inner  face,  with  a  prominent  basal  prostheca; 
mandibles  working  vertically  against  teeth  of  mentum.  Maxilla  (Plate  LXXXV,  458)  with 
palpi  very  short,  broad,  basal  segment  chitinized,  tip  narrow,  pale. 

Pupa. —  Length:  male,  10.4-11.4  mm.;  female,  10-13  mm. 
Width,  d.-s.;  male,  2-2.2  mm.;  female,  2.2-2.6  mm. 
Depth,  d.-v  ;  male,  1.9-2.2  mm.;  female,  2.1-2.5  mm. 

Living  pupae  with  pronotal  breathing  horns  light  yellow,  the  terminal  half  a  little  more 
brownish;  a  brownish  black  mark  on  prescutum;  abdomen  greenish,  more  yellow  behind; 
dorsal  spines  clear  light  green  thru  out  or  with  tips  in  some  specimens  a  little  infuscated. 
(In  pupae  preserved  in  alcohol,  the  greenish  colors  are  lost,  the  dark  brownish  black  mark 
of  the  mesonotal  prescutum  is  irregularly  U-shaped,  with  the  arms  of  the  U  directed  back- 
ward, and  the  dark  color  is  produced  caudad  and  cephalad  along  the  dorso-median  line; 
there  is  a  triangular  or  rounded  black  spot  on  either  side  of  the  scutellar  lobe;  on  the  metano- 
tum  is  a  large  blackish  median  blotch,  which  is  continued  cephalad  onto  the  mesonotal 
postnotum;  the  abdomen  has  an  interrupted  brownish  black  longitudinal  line  along  either 
side  of  the  midline  of  the  dorsum;  the  posterior  margin  of  each  tergite  is  suffused  with 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  973 

brown.)     In  old  and  fully  colored  pupae,  bases  of  dorsal  spines  brown,  with  the  tips  paler; 
head  and  thorax  with  appendages  brown,  in  some  specimens  very  dark;  abdomen  yellowish. 

Cephalic  part  of  head  very  flat  and  broad,  without  spines  but  with  a  small,  blunt  tubercle 
between  antennal  bases.  Labrum  transversely  wrinkled,  narrowed  to  the  bluntly  rounded 
apex.  Labial  lobes  large,  divergent,  each  lobe  rectangular  with  angles  rounded.  Sheaths* 
of  maxillary  palpi  rather  long,  curved  strongly  backward.  Antennal  sheaths  rather  enlarged, 
directed  cephalad,  bending  around  anterior  margin  of  eye  and  thence  directed  caudad, 
ending  just  beyond  knee  joint  of  fore  legs.  (In  older  pupae,  the  peculiar  nodose  antennal 
segments  of  the  adult  show  thru  the  sheath.)  Pronotal  breathing  horns  (Plate  LXXXV, 
464)  jlarge,  conspicuous,  directed  dorsad  and  laterad,  the  terminal  half  bent  rather 
suddenly  cephalad.  Mesonotum  transversely  wrinkled.  Metanotum  (Plate  LXXXV,  463) 
with  two  long,  slender  spines,  arising  beyond  midlength  of  segment,  directed  caudad  and 
slightly  dorsad.  Wing  sheaths  broad,  reaching  posterior  margin  of  second  abdominal  seg- 
ment. Leg  sheath  ending  just  before  posterior  margin  of  third  abdominal  segment;  fore  legs 
the  shortest,  hind  legs  the  longest. 

Abdominal  segments  with  a  narrow  basal  ring  and  a  broader  posterior  ring;  segment  1 
about  half  as  long  as  segment  2;  tergites  1  to  7  with  a  pair  of  long,  slender,  spinous  pro- 
jections, shortest  on  anterior  segment,  longest  on  seventh  segment,  these  projections  arising 
from  near  caudal  margin  of  segment,  directed  dorsad  and  caudad,  those  of  anterior  segment 
almost  parallel,  those  of  posterior  segments  more  divergent;  segments  2  to  7  having  lateral 
margins  produced  into  three  sharp  spines,  one  on  basal  ring  and  two  on  posterior  ring  of 
each  segment,  these  spines  directed  laterad  and  caudad,  the  terminal  spines  more  sharply 
caudad  than  the  other  two;  abdominal  sternites  armed  as  follows:  segment  3  with  a  small, 
subapical  spine  on  either  side,  these  spines  very  widely  separated,  segment  4  with  similar 
spines  but  larger  and  more  prominent,  segments  5  to  7  similarly  armed  but  with  another 
pair  of  small  spines  about  midlength  of  segment  and  much  nearer  midline  of  body,  segments 
2  to  7  with  a  subbasal  triangular  pit  on  either  side,  these  pits  widely  separated;  eighth 
tergite  with  caudal  margin  rounded,  concave,  the  lateral  angles  produced  backward,  upward, 
and  slightly  outward  as  strong  spines;  suture  on  ventral  surface  incomplete;  two  small  spines 
on  either  side  of  middle  line  of  body;  posterior  margin  of  segment  produced  caudad  as  two 
strong  spinous  projections.  Male  cauda  with  sternal  valves  rather  long,  tipped  with  two 
to  four  acute  spines,  in  some  specimens  with  two  spines  on  one  of  the  lobes  and  only  one 
on  the  other;  tergal  valves  a  little  rounded  at  tips,  slightly  longer  than  sternal  valves.  Female 
cauda  (Plate  LXXXV,  465  and  466)  with  sternal  valves  slender,  feebly  notched  at  tips; 
tergal  valves  broader,  rounded  at  tips,  and  with  a  deep  median  split. 

Nepionotype  —  Coy  Glen,  Ithaca,  New  York,  May  8,  1913. 
Neanotype  —  Ithaca,    May  30,  1913. 

Paratypes. —  Larvae  and  pupae  with  types;  others  from  Orono,  Maine,  June  17  and  19, 
1913. 

Genus  Triogma  Schiner  (Gr.  three  +  furrow) 

1863     Triogma  Schin.     Wien.  Ent.  Monatschr.,  vol.  7,  p.  223. 

Larva.' —  Body  covered  with  elongate  leaflike  projections,  some  of  the  dorsal  ones   with 
as  many  as  four  teeth  on  anterior  convex  face.     Spiracular  disk  surrounded  by  four  lobes. 
Pupa.—  Dorsum  of  abdomen  with  elongate  branched  spines. 


974  CHARLES  PAUL  ALEXANDER 

The  genus  Triogma  includes  but  three  known  species,  the  genotype 
Triogma  trisulcata  (Schum.)  of  Europe,  T.  kuwanai  (Alex.)  of  Japan, 
and  T.  exculpta  O.  S.  of  the  eastern  United  States.  The  last-named 
species  is  very  rare  and  its  habits  are  entirely  unknown. 

The  first  reference*  to  the  immature  stages  of  T.  trisulcata  is  by  Steinmann 
(1907-08),  who  discusses  the  larva  as  that  of  an  unknown  species  of 
Phalacrocera.  The  true  identity  of  this  insect  was  made  known  by 
Mliller  (1908-09).  Steinmann  found  the  larvae  at  Sackingen  at  the  end 
of  April,  fourteen  days  after  the  melting  of  the  snow.  They  were  found 
in  mountain  streams,  attached  to  and  living  among  the  stems  of  the 
aquatic  moss  Fontinalis  antipyretica  Linn.,  to  which  they  clung  firmly 
by  means  of  two  strong  chitinized  hooks  at  the  caudal  end  of  the  body. 
The  larva  resembles  to  a  startling  degree  the  moss  on  which  it  lives. 
Along  the  dorsal  surface  are  two  rows  of  leaflike  appendages,  each  of  the 
abdominal  segments  having  three  such  appendages,  of  which  the  most 
anterior  one  is  the  shortest  and  the  posterior  one  is  the  longest.  The 
anterior  appendage  is  untoothed,  the  others  have  as  many  as  four  teeth 
on  the  anterior  face.  The  pleural  region  likewise  bears  a  row  of  leaflike 
structures,  while  the  ventral  surface  shows  a  double  row  of  small,  knoblike 
leaflets.  Thru  the  strong  accentuation  of  the  dorsal  and  the  two  lateral 
rows,  there  is  produced  a  copy  of  the  ternate  condition  of  the  leaf  arrange- 
ment in  Fontinalis. 

The  larva  of  T.  trisulcata  (Plate  LXXXIII,  452)  is  of  a  light  green  color 
marked  with  darker  blotches,  and  measures  19  millimeters  in  length.  The 
longest  body  appendage  is  1.5  millimeters  in  length.  Miiller  found  larvae 
and  also  pupae.  The  pupa  is  characterized  by  the  possession  of  somewhat 
similar  appendages  to  those  of  the  larva,  the  dorsal  row  showing  the 
peculiar  branched  condition  found  in  the  larva. 

SUBFAMILY  Tipulinae 

The  subfamily  Tipulinae  comprises  a  remarkably  homogeneous  assem- 
blage of  usually  large  species.  It  is  made  up  of  a  relatively  few  but  in 
some  cases  very  extensive  genera,  which  are  found  thruout  the  world. 
Efforts  have  been  made  in  the  past  to  maintain  three  tribes  —  the 
Dolichopezini,  the  Ctenophorini,  and  the  Tipulini.  It  has  become  increas- 
ingly difficult,  however,  to  define  these  groups  on  the  constant  accession 
of  exotic  genera  and  species.  A  study  of  the  immature  stages  likewise 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  975 

i  .  i*«M     i,    i 

fails  to  substantiate  the  validity  of  these  groups,  and,  for  the  present 
at  least,  or  until  other  and  better  characters  may  be  found,  it  is  better 
to  eliminate  the  Dolichopezini  and  the  Ctenophorini,  founded,  as  they 
are,  on  insufficient  or  sexual  characters. 

The  species  of  the  Tipulinae  include  the  largest  crane-flies  known,  some 
exotic  species  of  Ctenacroscelis  attaining  a  wing  expanse  of  nearly  10 
centimeters.  In  North  America  the  largest  species  are  representatives 
of  the  genera  Longurio  and  Holorusia.  The  great  majority  of  the  species 
in  this  subfamily  are  well  above  the  average  in  size,  this  feature  alone 
being  sufficient  to  eliminate  all  but  a  very  few  of  the  other  groups  of 
Tipulidae.  The  smallest  member  of  the  subfamily  Tipulinae  known  to 
the  writer  is  Microtipula  amazonica  Alex.,  of  Brazil,  in  which  the  wing  of 
the  male  measures  but  7.2  millimeters  in  length  and  is  very  long  and 
narrow.  Practically  all  of  the  common  local  species  of  this  group  belong 
to  the  two  genera  Tipula  and  Nephrotoma. 

The  immature  stages  of  members  of  the  subfamily  Tipulidae  are  found 
in  a  wide  range  of  habitats.  Some  species  of  Tipula  are  almost  entirely 
aquatic  in  the  larval  state.  The  majority  of  the  known  species,  live  in 
moist  earth  near  water,  or  beneath  damp  cushions  of  moss.  Some,  such 
as  Oropeza,  live  in  much  drier  mosses  on  exposed  rocks.  A  rather  con- 
siderable number  of  species  (Brachypremna,  Ctenophora,  Dictenidia,  and 
several  species  of  Tipula)  live  in  decaying  wood  or  beneath  the  bark  of 
prostrate  trunks.  Tanyptera  lives  in  wood  which  is  relatively  sound, 
and  this  represents  the  extreme  development  of  this  tendency  in  the 
family. 

The  larvae  are  never  very  slender,  and  are  usually  very  plump  and 
terete.  In  a  few  cases  only  is  the  body  decidedly  depressed.  There  is 
a  definite  arrangement  of  setae  on  the  body,  there  being  none  on  the 
anterior  annulus  of  the  abdominal  segments  except  a  single  seta  on  the 
pleura  of  either  side.  The  spiracular  disk  is  surrounded  by  six  lobes, 
a  number  not  found  in  the  Limnobiinae.  In  Dolichopeza  the  number 
is  described  as  being  five,  the  normal  number  in  the  Eriopterini,  but 
all  other  features  of  the  genus  are  essentially  tipuline.  •  In  a  few  species 
the  number  of  lobes  is  increased  to  eight.  In  the  genus  Tanyptera  the 
lobes  are  exceedingly  reduced  in  size  so  that  the  caudal  end  appears  almost 
naked  and  exposed.  The  spiracles  are  always  present  and  in  some  cases 
are  very  large.  The  anal  gills  are  almost  always  present  and  are  variously 


976  CHARLES  PAUL  ALEXANDER 

developed  according  to  the  habitat.  The  head  capsule  is  remarkably 
uniform  thruout  the  group,  being  broad,  compact,  and  massive,  with  the 
posterior  incisions  shallow  and  the  prefrontal  sclerite  very  large  and 
usually  distinct.  The  labrum  is  usually  conspicuous,  transverse,  with 
fringes  of  hairs.  The  mentum  has  from  seven  to  nine  teeth  along  the 
anterior  margin,  and  is  deeply  split  behind  but  not  completely  divided. 
The  hypopharynx  consists  of  a  narrow,  flattened  plate,  with  the  basal 
lateral  angles  produced  into  strong  arms  and  the  anterior  margin  having 
usually  five  teeth,  the  teeth  being  rarely  more  numerous  and  in  some 
cases  obsolete.  The  antennae  are  cylindrical,  and  are  stoutest  in  the 
wood-inhabiting  species;  in  many  species  of  Tipula  and  Prionocera  they 
are  long  and  slender,  the  length  being  about  four  times  the  diameter; 
the  apical  papilla,  in  some  cases  obsolete,  is  usually  very  small  and  is 
flattened.  The  mandibles  are  not  large  in  proportion  to  the  size  of 
the  capsule;  they  have  few  teeth,  in  some  species  only  a  dorsal  and  a 
ventral  tooth  in  addition  to  the  apical  point;  the  prosthecal  appendage 
is  variously  developed.  The  maxillae  are  simple  and  generalized  in 
structure. 

The  pupae  are  fairly  uniform  in  structure  thruout  the  subfamily.  The 
tips  of  the  sheaths  of  the  maxillary  palpi  are  strongly  curved,  or,  in  the 
majority  of  species,  actually  recurved.  The  pronotal  breathing  horns  are 
variously  formed,  but  in  practically  all  species  they  are  short,  cylindrical, 
and  with  the  tips  but  little  expanded.  In  some  genera  (Longurio, 
Prionocera,  and  Tipulodina)  the  breathing  horns  are  greatly  elongated, 
and,  in  some  cases  at  least,  are  slightly  unequal  in  length,  the  longer 
measuring  nearly  half  the  length  of  the  body.  In  some  genera,  such 
as  Prionocera  and  Holorusia,  the  horns  are  split  at  their  tips  into  two 
divergent  flaps.  The  only  short,  clavate  horns  in  the  subfamily  are 
those  of  the  genus  Tanyptera.  The  mesonotum  is  often  provided  with 
four  or  six  variously  developed  tubercles.  The  abdominal  segments  are 
almost  always  armed  with  transverse  posterior  rows  of  spines,  these 
ranging  in  number  from  about  four  to  twenty. 

The  large  size  of  the  larvae  and  the  pupae  will,  as  a  rule,  easily  separate 
this  subfamily  from  almost  all  members  of  the  Limnobiinae.  The  few 
large  members  of  the  latter  group  are  readily  separated  by  the  characters 
outlined  above. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  977 

The  following  keys  separate  the  genera  of  the  subfamily  Tipulinae: 

Larvae  ^ ; 

1.  Spiracular  disk  surrounded  by  five  lobes;  living  in  moss Dolichopeza  Curt.  (p.  981) 

Spiracular  disk  surrounded  by  four  or  six  lobes,  or  with  lobes  indistinct ..--. 2 

2.  Spiracular  disk  with  lobes  indistinct;  living  in  nearly  solid  or  semi-decayed  wood. 

Tanyptera  Latr.  (p,  988) 
Spiracular  disk  with  lobes  distinct 3 

3.  Spiracular  disk  with  four  slender,  hornlike  lobes Tipula  (selene  Meig.)  (p.  1016) 

Spiracular  disk  with  six  lobes 4 

4.  Anal  gills  pinnately  branched.  .  . .Longurio  Loew  (p.  990) 

Aeshnasorna  Johns,  (p.  993) 
Anal  gills  not  pinnately  branched 5 

5.  Antepenultimate  segment  of  abdomen  with  a  strong  lateral  tubercle. 

Oropeza  Needm.  (p.  982) 
Antepenultimate  segment  of  abdomen  without  such  a  tubercle 6 

6.  Dorsum  of  head  behind  antenna  with  a  slender,  flexible  spine;  living  in  wood. 

Ctenophora  (apicata  O.  S.)  (p.  986) 
Dorsum  of  head  without  such  a  spine 7 

7.  Lobes  surrounding  spiracular  disk  elongate,  digitiform,  fringed  with  long  hairs. 

Prionocera  Loew  (p.  995) 
Lobes  surrounding  spiracular  disk  less  elongate 8 

8.  Size  large  (over  50  mm.);  spiracular  disk  with  the  six  moderately  long  lobes  fringed  with 

long  hairs;  mandible  small,  with  a  dorsal  and  a  ventral  tooth;  found  in  western  North 

America Holorusia  Loew  (p.  993) 

Size  smaller;  if  large  (T.  abdominalis) ,  the  lobes  surrounding  disk  bifid;  mandible  with 

two  or  three  ventral  teeth Tipula  Linn.  (p.  998) 

Nephrotoma  Meig.  (p.  1016) 
Pupae 

1.  Pronotal  breathing  horns  very  long,  slender,  the  longest  one  nearly  if  not  quite  half  length 

of  body 2 

Pronotal  breathing  horns  short,  cylindrical  or  flattened,  subequal  in  size 3 

2.  Length  40  mm.;  longest  breathing  horn  18  mm.;  maxillary  palpi  not  recurved  at  tips; 

venation  with  petiole  of  cell  Mi  very  short Longurio  Loew  (p.  990) 

Length  20  mm.;  longest  breathing  horn  9  to  10  mm.;  maxillary  palpi  recurved  at  tips; 
venation  with  petiole  of  cell  MI  longer Prionocera  Loew  (p.  995) 

3.  Pronotal  breathing  horns  short,  compressed,  deeply  bicrenulated;  living  in  wood. 

Tanyptera  Latr.  (p.  998) 
Pronotal  breathing  horns  cylindrical 4 

4.  Maxillary  palpi  not  recurved  at  tips 5 

Maxillary  palpi  recurved  at  tips • " 

5.  Pronotal   breathing   horns   with  apices   deeply  split;   mesonotum  with  two   tubercles; 

abdominal  segments  with  fourteen  to  thirty-four  spines;  found  in  Western  States; 

living  in  mud Holorusia  Loew  (p.  993) 

Pronotal  breathing  horns  short,  slender,  apices  not  split;  mesonotum  with  eight  tubercles; 
abdominal  segments  with  four  powerful  spines  near  posterior  margin;  found  in  Southern 
States;  living  in  wood Brachypremna  O.  S.  (p.  984) 

6.  Mesonotum  with  two  spines;  ventral  abdominal  spines  six  to  eight;  fore  and  middle  tarsi 

subequal,  shorter  than  hind  tarsi;  living  in  wood Ctenophora  Meig.  (p.  98o) 

Characters  not  as  above • •  •  •  •  • : 

7.  Mesonotum  with  a  large,  roughly  triangular,  reticulated  area  on  either  side  of  median 

line;  dorsum  of  cauda  with  four  lobes Oropeza  Needm.  (p.  982) 

Mesonotum  unarmed  or  with  four  or  six  lobes;  dorsum  of  cauda  with  six,  or  rarely  four 

I0bes Tipula  Linn.  (p.  998) 

Nephrotoma  Meig.  (p.  1016) 

10 


978  CHARLES  PAUL  ALEXANDER 

The  most  important  literature  on  the  Tipulinae  is  as  follows: 

Dolichopeza  albipes Pupa Beling,  1879:44^5  (as  sylvi- 

cola). 

Dolichopeza  albipes Larva,  pupa,  general . . .  Beling,  1886: 189-191  (as  syl- 

vicola). 

Tanyptera  atrata General Ferris,  1840:92;  1849:333. 

Tanyptera  atrata General Nordlinger,  1848. 

Tanyptera  atrata General De  Rossi,  1882. 

Tanyptera  atrata Larva,  pupa,  general . . .   Gercke,  1884,  PI.  I,  figs.  12-19. 

Tanyptera  atrata General Hermann,  1880. 

Tanyptera  atrata  ruficornis Larva Gerbig,    1913: 156-158    (as 

Ctenophora  flavicornis) . 

Tanyptera  fumipennis Larva,  pupa,  general . . .  Malloch,  1915-17  b:  194-195; 

1919. 

Dictenidia  bimaculata Larva Bouche*,  1834 : 32. 

Dictenidia  bimaculata General Zetterstedt,  1851 : 4023. 

Dictenidia  bimaculata Larva,  pupa,  general . . .   Weyenbergh,  1872. 

Dictenidia  bimaculata General Beling,  1873  b :  575. 

Dictenidia  bimaculata General Czizek,  1913: 102. 

Ctenophora  flaveolata Larva,  pupa,  general . . .   Reaumur,  1740,  PL  I,  fig.  9. 

Ctenophora  flaveolata General Zetterstedt,  185 1_:  4016. 

Ctenophora  flaveolata Larva,  pupa,  general . . .  Weyenbergh,  1872. 

Ctenophora  pectinicornis Larva,  pupa .' Bouch6,  1834: 29-31. 

Ctenophora  pectinicornis General Fischer  von  Waldheim,  1838. 

Ctenophora  pectinicornis Larva,  pupa Weyenbergh,  1872. 

Ctenophora  pectinicornis General Zetterstedt,  1851 : 4014. 

Ctenophora  pectinicornis General Kaltenbach,  1874: 202. 

Ctenophora  j 'estiva Larva Kaltenbach,  1874 : 631. 

Ctenophora  festiva Larva Czizek,  1911 : 48. 

Ctenophora  nigricoxa Pupa Lundstrom,  1906: 7. 

Ctenophora  apicata Larva,  pupa Johannsen,  1910:32-33. 

Ctenophora  angustipennis Larva,  pupa,  general. . .  Anthon,  1908. 

Ctenophora  angustipennis Egg,  larva,  pupa,  injury.  Lovett,  1915. 

Aeshnasoma  rivertonensis Larva Johnson,  1906: 1-2. 

Aeshnasoma  rivertonensis General Johnson,  1907-12  [1909]:  115- 

116. 

Tipulodina  pedata Pupa De  Meijere,  1911 : 64. 

Holorusia  rubiginosa Larva,  pupa,  general. . .   Kellogg,  1901,  a  and  b. 

Holorusia  rubiginoia Larva Comstock  and  Kellogg,  1904 : 

54-62. 

Prionocera  fuscipennis Larva,  pupa,  general. . .  Malloch,  1915-17b :  199-200. 

Prionocera  parri,  supp Larva Alexander,  1919  c. 

(Palaearctic  species) 

Tipula  caesia General Schiner,  1864 : 516. 

Tipula  dilatata Larva,  pupa Beling,  1886: 176. 

Tipula  dilatata Larva,  pupa Czizek,  1913: 169. 

Tipula flavolineata General Staeger,  1840:23. 

Tipula  flavolineata Larva,  pupa Beling,  1873  b :  581-582. 

Tipula  flavolineata Larva,  pupa Czizek,  1913 : 146. 

Tipula  fulvipennis Larva,  pupa Beling,   1879: 22-24   (as  lutes- 

cens). 

Tipula  fulvipennis Larva,  pupa Czizek,  1913: 74. 

Tipula  hortensis Larva,  pupa Beling,  1873  b:  578-579. 

Tipula  hortensis Larva Gerbig,  1913: 154-156. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  979 

Tipula  hortulana Larva,  pupa,  general . . .   Beling,  1879:25;  1886: 178- 

179. 

Tipula  hortulana Larva,  pupa,  general . . .  Czizek,  1913:96. 

Tipula  imbecilla General Loew,  1869: 9. 

Tipula  irrorata"1 Larva,  pupa Beling,  1873  b:  586-587. 

Tipula  irrorata Larva,  pupa Czizek,  1913:98-99. 

Tipula  lateralis Larva,  pupa Beling,  1879: 26-28. 

Tipula  lateralis Larva Gerbig,  1913: 153-154. 

Tipula  lateralis Larva,  pupa Czizek,  1913: 128. 

Tipula  lateralis General Cameron,  1917:61.]      / 

Tipula  lunata Larva,  pupa Brocher,  1909. 

Tipula  luteipennis Larva,  pupa Beling,  1886: 181-182. 

Tipula  luteipennis Larva,  pupa Czizek,  1913: 143. 

Tipula  marmorata Larva,  pupa . .  Beling,  1886: 182-183. 

Tipula  maxima Larva,  pupa Beling,  1886: 177-178  (as  gigan- 

tea). 

Tipula  maxima Larva Gerbig,  1913: 152-153  (as gigan- 

tea). 

Tipula  maxima Larva,  pupa Czizek,  1913 : 70-71. 

Tipula  maxima General Wesenberg-Lund,  1915:335  (as 

gigantea). 

Tipula  micans1 Larva,  pupa Beling,  1886: 183-184. 

Tipula  nigra Larva,  pupa Beling,  1879 : 28-29. 

Tipula  nigra Larva,  pupa Czizek,  1913: 132. 

Tipula  nubeculosa Larva,  pupa Beling,  1873  b :  575-577. 

Tipula  nubeculosa Larva,  pupa Czizek,  1913: 113. 

Tipula  ochracea Larva,  pupa Beling,  1873  b :  582-583. 

Tipula  ochracea Larva,  pupa Czizek,  1913 : 157. 

Tipula  oleracea Larva,  pupa Del  Guercio,  1914. 

Tipula  oleracea General Patterson,  1908. 

Tipula  pabulina. Larva,  pupa Beling,  1873  b: 579-580. 

Tipula  pabulina. Larva,  pupa. Czizek,- 1913:86. 

Tipula  pagana Larva,  pupa Beling,  1879: 29-31. 

Tipula  paludosa .  .  Larva,  pupa Beling,  1873  b :  583-585. 

Tipula  paludosa Larva Gerbig,  1913:136-151. 

Tipula  paludosa Larva,  pupa Czizek,  1913: 136. 

Tipula  paludosa Larva,  pupa Rennie,  1916;  1917. 

Tipula  parva,  supp General Onuki,  1905. 

Tipula  peliostigma .  . Larva,  pupa Beling,  1879 : 33-34. 

Tipula  peliostigma ! Larva,  pupa Czizek,  1913: 159. 

Tipula  pruinosa Larva,  pupa Beling,  1879: 31-32;  1886: 184. 

Tipula  pruinosa Larva,  pupa Czizek,  1913: 148. 

Tipula  rufina Larva,  pupa Mik,  1882  a. 

Tipula  scripta ^ Larva,  pupa Beling,  1873  b: 577-578. 

Tipula  scripta f Larva,  pupa Czizek,  1913: 110. 

Tipula  selene Larva,  pupa Beling,  1879: 34-35. 

Tipula  selene Larva,  pupa Czizek,  1913: 160-161. 

Tipula  signata Larva,  pupa Beling,  1879: 32-33. 

Tipula  signata Larva,  pupa Czizek,  1913: 104. 

Tipula  simplex General Doane,  1908. 

Tipula  subnodicornis Larva,  pupa Beling,  1886: 185-186. 

Tipula  truncorum Larva,  pupa Beling,  1879: 24-25. 


^  Tipula  micans  is  considered  by  Kertesz  and  others  as  a  synonym  of  T.  irrorata,  but  Beling's  descrip- 
tions indicate  that  he  had  two  distinct  species  before  him. 


980  CHARLES  PAUL  ALEXANDER 

Tipula  truncorum Larva,  pupa Czizek,  1913:82. 

Tipula  unca Larva,  pupa,  general . . .   Beling,  1886: 179-181  (as  longi- 

cornis). 
Tipula  unca Larva,  pupa Czizek,  1913: 101-102  (as  longi- 

cornis). 
Tipula  variicornis Larva,  pupa Beling,  1886: 173-174  (as  annu- 

licornis). 

Tipula  variicornis Larva,  pupa Czizek,  1913: 122-123. 

Tipula  variipennis Pupa Beling,  1873  b:580-581. 

Tipula  variipennis Larva Beling,  1886: 186. 

Tipula  variipennis General Westhoff ,  1879. 

Tipula  variipennis Larva Gerbig,  1913: 131-136. 

Tipula  variipennis Larva,  pupa Czizek,  1913:91-92. 

Tipula  vernalis Larva,  pupa Beling,  1879:25-26. 

Tipula  vernalis Larva,  pupa Czizek,  1913: 120. 

Tipula  vittata Larva,  pupa Beling,  1886: 186-188. 

Tipula  vittata. . Larva,  pupa Czizek,  1913 : 78. 

Tipula  winnertzii* Larva,  pupa Beling,  1873  b:  585-586;  1886: 

188-189. 

Tipula  winnertzii Larva,  pupa Czizek,  1913:84-85. 

(Nearctic  species) 
Tipula  abdominalis Larva Malloch,  1915-17  b: 200-201  (as 

Tipula  sp.  2). 

Tipula  arctica Larva,  pupa Nielsen,  1910: 57-59. 

Tipula  arctica Larva,  pupa Alexander,  1919  c:  18c,  19c. 

Tipula  bicornis Larva,  pupa Forbes,  1890. 

Tipula  caloptera Larva Needham  and  Betten,  1901 : 

575-576  (as  abdominalis). 

Tipula  cunctans Larva,  pupa,  general .  . .  Hyslop,  1910  (as  infuscata). 

Tipula  cunctans. Larva,  pupa Malloch,  1915-17 b: 204. 

Tipula  eluta Larva,  pupa,  general. .  .   Hart,  1898  [1895]:  212-214. 

Tipula  eluta Larva,  pupa Malloch,  1915-17  b:203. 

Tipula  serta  (?) Pupa Malloch,  1915-17  b:205. 

Tipula  trivittata Larva,  pupa Greene,  1909. 

Tipula  trivittata Pupa Malloch,  1915-17b:  204-205. 

Tipula  ultima Pupa,  general Needham,  1903:280-281  (as 

flavicans). 

Tipula  ultima General Caudell,  1913  (as  flavicans). 

Habromastix  cinerascens General Skuse,  1890 : 95. 

Nephrotoma  analis Larva,  pupa Beling,  1886: 172-173. 

Nephr-otoma  analis Larva,  pupa Czizek,  1911 : 70-71. 

Nephrotoma  cornicina Larva,  pupa Beling,  1879 : 39-40  (as  iridi- 

color). 

Nephrotoma  cornicina Larva,  pupa Czizek,  1911:76. 

Nephrotoma  crocata Larva,  pupa Beling,  1879:40-41. 

Nephrotoma  crocata Larva,  pupa Czizek,  1911 : 87. 

Nephrotoma  lineata Larva,  pupa Beling,  1879:42-43  (as  histrio). 

Nephrotoma  lineata Larva,  pupa Czizek,  1911 : 83-84. 

Nephrotoma  lunulicornis Larva,  pupa Beling,  1879:41-42. 

Nephrotoma  lunulicornis Larva,  pupa Czizek,  1911:61. 

Nephrotoma  maculata Larva,  pupa Beling,  1879: 36-37. 

Nephrotoma  maculata Larva,  pupa Czizek,  1911 : 80-81. 

Nephrotoma  pratensis Larva,  pupa Beling,  1886: 175-176. 

8Riedel  (1913:25)  considers  Tipula  winnertzii  as  a  synonym  of  T.  truncorum,  but  Beling's  descriptions 
indicate  that  the  larvae  he  had  before  him  represented  two  distinct  species. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  981 

Nephrotoma  pratensis Larva,  pupa Czizek,  1911 : 85-86. 

Nephrotoma  quadrifaria Larva,  pupa Beling,  1879:37-39. 

Nephrotoma  quadrifaria Larva,  pupa Czizek,  1911:66. 

Nephrotoma  ferruginea Larva,  pupa Hart,  1898  [1895] :  218-219. 

Nephrotoma  ferruginea Larva,  pupa Malloch,  1915-17  b:206. 

Tribe  Tipulini 
Subtribe  Dolichopezaria 
Genus  Dolichopeza  Curtis  (Gr.   long  +feet) 

1825     Dolichopeza  Curt.     Brit.  Ent.,  p.  62. 

1830    Leptina  Meig.     Syst.  Beschr.  Zweifl.  Ins.,  vol.  6,  pi.  65,  fig.  10. 

1846    Apeilesis  Macq.     Dipt.  Exot.,  Suppl.  1,  p.  8. 

The  genus  Dolichopeza  is  a  small  group  of  flies  including  about  a  score 
of  species,  most  numerous  in  the  Oriental  and  Australian  regions.  The 
immature  stages  of  the  European  Dolichopeza  albipes  Strom,  a  species 
that  is  very  close  to  the  American  species  D.  americana  Needm.,  have 
been  described  by  Beling  (1879,  1886).  He  found  larvae  in  and  beneath 
moss  cushions  covering  the  piles  of  waste  copper  slag  in  the  Harz  Moun- 
tains. On  June  11,  1878,  larval  and  pupal  material  was  taken  from  a 
tussock  of  the  Jungermanniales  liverwort  Alicularia  scalaris  Corda.  The 
pupal  period  was  found  to  be  six  days.  Females  were  noted  depositing  their 
eggs  in  these  hummocks,  and  small  swarms  of  males  were  observed  dancing 
near  by.  It  appears  that  the  larvae  feed  on  the  upper  side  of  the  moss 
cushion  at  night,  withdrawing  into  the  interior  at  other  times.  A  larva  was 
found  in  earth,  which  shows  that  these  larvae  are  not  dependent  on  mosses. 

The  larva  is  described  by  Beling  (1886:189-191)  as  being  about  12 
millimeters  long  and  2.3  millimeters  in  diameter.  The  body  is  almost 
terete.  The  color  is  a  rather  bright  green,  the  dorsum  being  marked 
with  two  zigzag  dark  brown  stripes.  The  spiracular  disk  has  but  five 
lobes;  the  three  dorsal  lobes  are  long  and  narrow,  the  median  tooth  being 
formed  by  the  apparent  fusion  of  two  teeth;  the  ventral  lobes  are  very 
short,  are  tuberculate,  and  have  a  small,  dark  brown,  triangular  mark 
at  the  inner  tip.  There  is  a  small  dark  cross-stripe  at  the  base  of  each 
lateral  lobe.  The  spiracles  are  small,  are  circular,  and  are  separated  by 
a  distance  equal  to  about  one  and  one-half  the  diameter  of  one.  Beneath 
each  spiracle  is  a  rather  large,  irregular,  blackish  brown  spot.  The  anal 
gills  are  strongly  protuberant.  The  pupa  measures  about  14.5  millimeters 
in  length  and  2.5  millimeters  in  diameter.  The  pronotal  breathing  horns 


982  CHARLES  PAUL  ALEXANDER 

are  directed  straight  outward  and  finally  downward.  The  eighth  segment 
has  a  circle  of  four  dorsal,  four  ventral,  and  two  pleural  spines,  which 
are  finely  bifid  at  their  tips.  The  pupa  is  green,  as  is  the  larva,  with  similar 
angular  dorsal  stripes. 

This  is  the  only  tipuline  larva  known  to  the  writer  with  five  lobes  sur- 
rounding the  spiracular  disk.  This  feature  is  the  more  remarkable  when 
this  larva  is  compared  with  that  of  Oropeza,  apparently  closely  related 
but  with  a  normal  tipuline  appearance.  The  immature  stages  of  other 
species  of  Dolichopeza  and  related  genera  will  be  of  interest. 

The  resting  positions  of  Dolichopeza  are  described  on  page  713  of  this 
paper,  and  the  striking  dissimilarities  to  Oropeza  noted.  It  may  be  that 
Oropeza  is  not  so  close  to  Dolichopeza  as  has  been  believed.  Osten  Sacken 
(1886:157)  describes  the  mating  of  a  European  Dolichopeza,  presumably 
D.  albipes,  as  follows: 

I  had  occasion  to  observe  the  copula  (in  Heidelberg,  July  26) ;  the  female  was  hanging  down 
from  some  support  to  which  it  held  on  by  its  front  legs;  it  bore  the  whole  weight  of  the  male, 
who  was  fastened  to  it  merely  by  the  forceps,  hanging  head  downwards,  with  his  legs  stretched 
out.  I  have  seen  Bittacomorpha  copulate  in  the  same  manner. 

Genus  Oropeza  Needham  (Gr.  mountain  +  feet) 

1908    Oropeza  Needm.     23d  Kept.  N.  Y.  State  Ent.  (1907),  p.  211. 

Larva. —  Form  somewhat  depressed.  A  strong  tubercle  on  sides  of  body  before  spiracular 
disk.  Spiracular  disk  surrounded  by  six  lobes,  the  four  dorsal  ones  slender,  the  ventral 
pair  blunt.  Anal  gills  blunt.  Head  capsule  of  the  tipuline  type.  Mandible  blunt,  with  about 
seven  teeth.  Hypopharynx  five-toothed.  Mentum  seven-toothed.  Coloration  dark  green. 

Pupa. —  Pronotal  breathing  horns  elongate-cylindrical,  slender.  Mesonotum  with  a 
slightly  elevated  triangular  area  on  either  side  of  mid-dorsal  line.  Leg  sheaths  ending  about 
on  a  level,  or  fore  legs  a  little  longer.  Abdominal  segments  with  two  stout  pleural  spines; 
posterior  annul!  before  margin  with  a  transverse  row  of  twelve  or  fewer  spines. 

Oropeza  is  a  genus  of  crane-flies  including  ten  described  species,  all 
of  which  occur  in  eastern  North  America  with  the  exception  of  three 
Japanese  forms.  The  flies  are  common  beneath  dark  culverts  and  bridges, 
in  outhouses,  in  crannies  and  crevices  of  rocky  cliffs,  beneath  overhanging 
boulders  along  mountain  streams,  and  in  similar  situations.  The  resting 
positions  of  the  adult  flies  are  discussed  on  page  712.  Many  species  show 
a  notable  predilection  for  resting  on  spiders'  webs.  The  immature  stages 
of  Oropeza  obscura  are  spent  in  dry  moss,  as  discussed  below.  Other 
species  have  been  reared  in  sandy  soil  and  in  wet  moss. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  983 

Oropeza  obscura  Johns. 

1909  Oropeza  obscura  Johns.  Proc.  Boston  Soc.  Nat.  Hist.,  vol.  34,  p.  122. 
Mr.  Hyslop  has  sent  to  the  writer,  at  various  times  during  the  past 
two  or  three  years,  larvae  of  Oropeza  obscura.  Numerous  larvae  and 
one  pupal  skin  were  found  under  dry  moss  (Hedwigia  albicans  [Web.] 
Lindb.)  on  rocks  in  the  South  Mountains,  Maryland,  on  November  4, 
1916,  by  H.  L.  Parker.  They  were  associated  with  the  larvae  of  a  dascillid 
beetle,  Eurypogon  niger  (Melsh.).  Additional  specimens  were  found  in 
a  decaying  log,  but  the  preferred  habitat  seems  to  be  beneath  moss.  The 
dark  green  larvae  are  very  sluggish. 

Larva. —  Length,  11-12  mm. 

Diameter,  1.7-1.8  mm. 

Color  dark  brownish  green,  ventral  surface  clearer  green. 

Form  depressed;  body  short  and  stout.  Dorsum  covered  with  small,  blackened  points, 
producing  the  dark  color  of  this  region. of  the  body.  Segments  much  wrinkled,  especially 
on  basal  ring.  On  each  side  of  penultimate  segment  a  stout  tubercle  which  simulates  dorsal 
lobes  of  spiracular  disk.  Spiracular  disk  (Plate  LXXXVI,  469)  moderate  in  size,  surrounded 
by  six  fleshy  lobes,  dorsal  pair  short  and  slender,  lateral  pair  long  and  slender,  ventral  pair 
very  short  and  blunt;  ventral  lobes  sparsely  fringed  with  short,  black  hairs,  and  with  a  pale 
apical  area  bearing  a  sensory  bristle;  inner  face  broadly  suffused  with  brown;  at  base  of 
other  lobes  similar  but  smaller  triangular  brown  marks;  a  dusky  area  ventrad  of  spiracles. 
Spiracles  large,  placed  obliquely.  Anal  gills  four,  large  and  blunt,  pale  yellow. 

Head  capsule  of  the  massive  tipuline  type.  Labrum  broad,  with  a  short,  blunt  lobe  on 
either  side;  median  epipharyngeal  region  with  dense  rows  of  stout  setae  and  two  small 
papillae  on  margin;  lateral  lobes  with  about  four  or  five  sensory  bristles  or  papillae  of  various 
sizes.  Mentum  (Plate  LXXXVI,  467)  seven-toothed,  median  point  long  and  slender,  a  pro- 
longation of  outer  mental  plate.  Hypopharynx  (Plate  LXXXVI,  468)  five-toothed,  the 
three  intermediate  teeth  the  largest,  lateral  tooth  on  either  side  small.  Antenna  slender, 
cylindrical,  with  apical  segment  very  reduced.  Mandible  rather  slender,  with  about  seven 
blunt  teeth  on  margin.  Maxilla  with  cardq  large,  triangular,  with  two  setifero us  punctures, 
the  outermost  very  large  and  hyaline  and  with  two  long,  powerful  setae;  lobes  of  maxilla 
fringed  with  dense  hairs;  palpus  short,  disklike,  with  a  large  group  of  tiny  hyaline  papillae 
at  apex. 

Pupa. —  Length  of  cast  skin,  about  12  mm. 

Coloration  brown;  pronotal  breathing  horns  dark  blackish  brown.  (It  is  possible  that 
the  pupae  when  alive  are  dark  green  in  color,  like  the  larvae.) 

Labrum  triangular.  Labial  lobes  large,  separated  by  apex  of  labrum.  Sheaths  of  maxillary 
palpi  short  and  stout,  apex  recurved  to  beyond  midlength.  Antennal  sheaths  moderately 
elongated.  Pronotal  breathing  horns  elongate,  slender,  cylindrical,  tips  a  little  expanded. 
Mesonotum  on  either  side  of  median  line  with  a  large,  roughly  triangular,  reticulated  area 
which  is  slightly  elevated,  with  apex  of  triangle  directed  toward  mid-dorsal  line.  Mesonotum 


984  CHARLES  PAUL  ALEXANDER 

transversely  wrinkled.  Metanotum  (Plate  LXXXVI,  470)  near  anterior  margin  with  a  trans- 
verse row  of  setiierous  punctures,  there  being  four  on  either  side,  the  intermediate  ones  with 
two  setae.  Wing  sheaths  reaching  base  of  third  abdominal  segment.  Leg  sheaths  reaching 
beyond  midlength  of  fourth  abdominal  segment,  fore  legs  a  little  the  longest,  hind  legs 
a  little  the  shortest. 

First  abdominal  segment  on  dorsum  somewhat  similar  to  metanotum,  with  a  transverse 
row  of  four  punctures  at  about  midlength,  the  lateral  ones  with  two  setae;  remaining 
abdominal  segments  (Plate  LXXXVI,  471)  subdivided  into  approximately  equal  basal  and 
posterior  rings;  on  tergites,  posterior  ring  with  a  transverse  row  of  spines,  basal  ring  unarmed; 
on  sixth  tergite,  where  best  developed,  spines  on  posterior  ring  numbering  about  twelve,  with 
a  seta  at  each  end  of  the  row  and  two  others  interspersed;  on  anterior  tergites,  spines  very 
email;  on  seventh  tergite  spines  large,  elongated;  on  eighth  tergite  four  powerful  spines  in 
transverse  alinement;  on  pleurites,  a  strong  spine  on  each  ring,  that  of  basal  ring  deeply 
bifid  and  with  a  strong  seta  in  notch  thus  formed;  posterior  ring  with  a  single  spine  bearing 
a  long  seta  on  its  face;  on  sternites,  condition  generally  similar  to  that  on  tergites,  the  spines 
a  little  more  prominent  but  of  about  the  same  number,  these  spines  slender  at  their  slightly 
curved  tips  and  lacking  on  sternites  2  to  4.  Female  cauda  (Plate  LXXXVI,  472  and  473) 
with  acidothecae  elongate,  contiguous,  on  either  side  of  tergites  at  base  with  a  powerful 
incurved  hook;  posterior  margin  of  eighth  tergite  with  four  spines,  lateral  pair  a  little  the 
larger;  ventral  side  at  base  with  a  median  protuberance  bearing  two  powerful  lateral  lobes 
and  two  smaller  chitinized  spines  which  are  directed  caudad  and  ventrad;  base  of  segment 
8  with. a  crossrow  of  eight  spines;  posterior  ring  of  segment  8  with  a  powerful  spine  at 
lateral  end. 

Nepionotype. —  South  Mountains,  near  Myersville,  Maryland,  March  31,  1916. 
Neanotype. —  Hagerstown,   Maryland,   May  9,    1916;   cast  pupal  skin  in  collection  of 
United  States  National  Museum. 

Paratypes. —  Larvae  from  type  locality. 

Genus  Brachypremna  Osten  Sacken  (Gr.  short  +  trunk) 
1886    Brachypremna  0.  S.    Berl.  Ent.  Ztschr.,  vol.  30,  p.  161. 

Larva. —  Unknown. 

Pupa. —  Antennal  segments  enlarged  at  base.  Antenna  short.  Sheaths  of  maxillary 
palpi  short,  not  recurved  at  tips.  Pronotal  breathing  horns  short,  slender,  finely  annulated. 
Mesonotum  with  eight  prominent  tubercles.  Leg  sheaths  reaching  beyond  midlength  of 
fourth  abdominal  segment,  fore  tarsi  very  short,  other  tarsi  subequal  in  length.  Abdominal 
segments  each  with  four  slender  spines  on  posterior  ring  of  both  sternites  and  tergites,  just 
before  posterior  margin;  two  pleural  spines;  eighth  segment  of  male  with  four  prominent, 
spinous  lobes. 

Brachypremna  is  a  small  genus,  including  but  nine  recent  species  found 
in  the  Austral  and  Tropical  regions  of  the  New  World.  Brachypremna 
eocenica  Meunier  is  described  from  the  Baltic  amber.  The  flies  of  the 
best-known  species,  B.  dispellens,  are  known  in  parts  of  the  Southern 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  985 

States  as  "  weavers."  They  frequent  rather  shady  places  and  have 
a  remarkable  dance  over  three  or  four  feet  of  vertical  space,  whence  the 
name  "  king  of  the  dancing  tipulids  "  given  them  by  Johnson.  This 
species  is  the  only  one  whose  immature  stages  are  at  all  known. 

Brachypremna  dispellens  (Walk.) 

1860     Tipula  dispellens  Walk.     Trans.  Ent.  Soc.  London,  n.  ser.,  vol.  5,  p.  333-334. 
1886    Brachypremna  dispellens  O.  S.     Berl.  Ent.  Ztschr.,  vol.  30,  p.  162. 

Brachypremna  dispellens  is  the  most  widely  distributed  species  of  the 
genus.  It  ranges  from  New  Jersey  southward  thru  North  America,  and 
thru  South  America  as  far  as  Argentina.  A  larva  found  by  R.  C.  Shannon 
in  a  rotten  log  by  a  stream  near  Washington,  D.  C.,  on  April  23,  1913, 
was  placed  in  rearing  and  emerged  in  May  as  an  adult  male  of  this  species. 
The  badly  mutilated  pupal  skin  was  preserved  and  is  here  described. 
No  part  of  the  larva  was  preserved. 

Pupa. —  Length  of  cast  pupal  skin,  about  18  mm. 

Coloration  brownish  yellow;  abdomen  with  a  broad  brown  sublateral  stripe  on  both  ventral 
and  dorsal  segments;  each  of  pleural  spines  set  in  a  brown  spot. 

Head  small.  Antennal  spines  very  large  and  crowded  at  base,  soon  passing  into  the  short, 
slender  flagellum.  Labrum  (Plate  LXXXVII,  474)  blunt.  Labial  lobes  closely  approximated, 
so  as  to  appear  as  a  single  large,  transversely  rectangular  lobe  at  end  of  labrum.  Sheaths 
of  maxillary  palpi  short,  not  recurved  at  tip.  Pronotal  breathing  horn  (Plate  LXXXVII,  475) 
small,  slender,  curved,  ringed  with  fine  annuli,  tapering  gradually  to  the  small  apex;  margin 
of  apex  set  with  breathing  pores.  Mesonotum  with  eight  conspicuous,  blunt,  naked  tubercles; 
the  four  intermediate  tubercles  larger,  arranged  in  the  form  of  a  trapezoid;  anterior  median 
pair  high,  conical,  located  rather  close  to  mid-dorsal  line.  Wing  sheaths  reaching  end  of 
second  abdominal  segment.  Leg  sheaths  (Plate  LXXXVII,  476)  extending  beyond  mid- 
length  of  fourth  abdominal  segment;  fore  legs  very  short,  ending  opposite  base  of  third 
tarsal  segment  of  other  legs. 

Abdominal  tergite  1  with  a  pair  of  long,  slender  spines  before  posterior  margin;  segments  2 
to  7  subdivided  into  a  basal  and  a  posterior  ring,  the  latter  with  a  transverse  row  of  four  long, 
slender  spines  before  posterior  margin,  the  seventh  tergite  with  about  six  such  spines;  sternites 
similar,  with  four  spines  on  posterior  ring;  pleurites  with  a  slender  spine  on  basal  and  posterior 
ring;  at  base  of  posterior  ring  between  spines,  an  indistinct,  slightly  protuberant  spiracle. 
Male  cauda  (Plate  LXXXVII,  477)  narrowed,  small,  valves  blunt;  on  dorsal  side  near  base 
four  conspicuous  lobes,  each  terminating  in  a  slender,  chitinized  spine;  a  small  acute  spine  on 
sides  of  ninth  segment  at  base. 

Neanotype.—  Cast  pupal  skin,  Washington,  D.  C.,  May,  1913  (in  collection  of  United 
States  National  Museum). 


CHARLES  PAUL  ALEXANDER 

Subtribe  Ctenophoraria 

Genus  Ctenophora  Meigen  (Gr.  comb  +  to  bear) 

1800    Flabellifera  Meig.     Nouv.  Class.  Mouch.,  p.  13  (nomen  nudum). 

1803     Ctenophora  Meig.     Illiger's  Mag.,  vol.  2,  p.  263. 

1910     Phoroctenia  Coq.     Proc.  U.  S.  Nat.  Mus.,  vol.  37,  p.  589. 

Larva. —  Body  opaque,  integument  rather  thick.     Spiracular  disk  surrounded  by  six  lobes. 

Pupa. —  Sheaths  of  maxillary  palpi  recurved.  Pronotal  breathing  horns  long  and  slender 
Two  spines  on  mesonotum.  Fore  and  middle  tarsi  subequal,  a  little  shorter  than  hind 
tarsi.  Ventral  abdominal  segments  with  six  to  eight  spines. 

Ctenophora  is  a  small  genus  including  about  sixteen  nominal  species 
found  thruout  the  Holarctic  region.  The  larvae  and  the  pupae  occur 
in  decaying  wood.  The  early  stages  have  long  been  known,  having  been 
described  by  Reaumur  and  Ds  Geer. 

In  Europe,  Ctenophora  flaveolata  (Fabr.)  is  described  and  figured  by 
Reaumur  (1740)  and  by  Weyenbergh  (1872).  C.  pectinicornis  (Linn.) 
is  described  or  mentioned  by  Bouche*  (1834),  by  Fischer  von  Waldheim 
(1838),  by  Zetterstedt  (1851:4014),  by  Weyenbergh  (1872),  and  by 
Kaltenbach  (1874).  C.  f estiva  Meig.  was  reared  by  Kaltenbach  (1874:631) 
from  larvae  in  decayed  beech  stems.  C.  nigricoxa  Lundst.  (Malpighia 
vittata  Meig.,  auct.  Frey)  was  reared  by  Lundstrom  (1906:7)  from  pupae 
in  rotten  birch  stumps.  The  immature  stages  of  the  various  European 
species  of  Ctenophora  are  described  as  living  in  the  wood  of  various 
trees  such  as  willow  (Salix),  birch  (Betula),  cherry  (Prunus),  and  other 
hardwood  species. 

In  North  America,  C.  apicata  is  described  by  Johannsen  (1910)  from 
elm  (Ulmus),  and  C.  angustipennis  Loew  by  Anthon  (1908)  in  alder  (Alnus) 
and  in  poplar  (Populus),  The  latter  species  is  recorded  also  as  injuring 
prune  trees  (Prunus)  in  Oregon,  by  Lovett  (1915),  who  gives  an  excellent 
account  of  all  stages  and  the  type  of  injury  done.  The  female  lays  from 
200  to  400  eggs,  which  hatch  in  from  nine  to  seventeen  days  and  the 
larvae  tunnel  into  the  surrounding  dead  wood.  Here  they  feed  and  grow, 
reaching  maturity  the  following  spring.  Pupation  takes  place  in  the 
burrows,  the  pupal  stage  requiring  about  ten  days.  Osten  Sacken 
(1877:211)  supposed  that  the  larvae  live  in  the  stumps  of  redwood 
(Sequoia),  but  this  has  never  been  confirmed. 

Ctenophora  apicata  O.  S. 

1864     Ctenophora  apicata  O.  S.     Proc.  Ent.  Soc.  Phila.,  vol.  3,  p.  46. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  987 

The  larvae  and  the  pupae  of  Ctenophora  apicata  that  were  described 
by  Johannsen  (1910)  have  been  studied  by  the  writer  in  the  collection 
of  the  Maine  Agricultural  Experiment  Station.  They  were  collected  at 
Orono,  Maine,  on  June  23,  1909,  by  Dr.  William  C.  Woods.  In  1913 
the  writer  examined  the  stump  from  which  they  were  taken,  and  found 
a  few  cast  pupal  skins. 

The  notes  here  given  are  taken  partly  from  Dr.  Johannsen's  description 
and  partly  from  the  original  material. 

Larva. —  Length,  about  30  mm. 

Color  white.  Body  stout,  cylindrical.  Antenna  cylindrical,  with  an  apical  paoilla. 
On  dorsum  of  head  behind  antennae,  a  slender,  flexible  spine.  Spiracular  disk  surrounded 
by  six  lobes,  dorsal  and  lateral  pairs  slender,  finger-like;  ventral  pair  blunt. 

Pupa. —  Length,  25-27  mm. 

Length  of  breathing  horns,  3  mm.  additional. 
Width,  d.-s.,  5.2  mm. 
Depth,  d.-v.,  5  mm. 

Pupa  somewhat  similar  to  that  of  Tanyptera,  differing  as  follows:  Form  stout;  abdomen 
a  little  depressed.  Sheaths  of  maxillary  palpi  strongly  recurved  at  tips.  Pronotal  breathing 
horns  long  and  slender.  Mesonotum  with  a  prominent  tuberculate  spine  on  either  side  of 
median  line.  Legs  shcrt,  ending  before  tip  of  third  abdominal  segment,  tarsal  sheaths  ending 
about  on  a  level,  or  those  of  hind  legs  a  very  little  longer.  Abdominal  segments  5  to  7  with 
six  to  eight  spines.  Female  cauda  with  six  powerful  ventral  spines  or  tubercles  and  two 
dorsal  tubercles  on  either  side.  Valves  of  ovipositor  short,  tergal  valves  a  little  longer  than 
sternal  valves. 

Nepionotype. —  Orono,  Maine,  July,  1909. 

Neanotype. —  Orono,  Maine,  July,  1909. 

Paratypes. —  Numerous  pupae   in  collection  of  Maine  Agricultural  Experiment  Station. 

Genus  Dictenidia  Brulle  (Gr.   double  +  comb) 

1833     Dictenidia  Brulle.     Ann.  Soc.  Ent.  France,  vol.  2,  p.  401-402. 

1856     Ceroctena  Rond.     Dipt.  Ital.  Prodr.,  vol.  1,  p.  186. 

1863    Dicera  Lioy.     Atti  dell'  Institut  Veneto,  ser.  3,  vol.  9,  p.  216. 

Dictenidia  is  a  genus  of  Palaearctic  crane-flies  including  three  species, 
of  which  one  is  European  and  the  others  are  Japanese.  The  genotype, 
Dittenidia  bimaculata  Brulte,  is  very  well  known.  The  immature  stages 
are  described  or  mentioned  by  Bouch4  (1834),  by  Zetterstedt  (1851),  by 
Weyenbergh  (1872),  by  Beling  (1873  b),  by  Czizek  (1913),  and  by  other 
investigators.  Beling  found  the  larvae  in  decaying  birch  (Betula).  He 
describes  the  larvae  as  being  grayish  yellow,  with  four  spiracular  lobes. 
The  pupal  duration  is  seven  days.  Osten  Sacken  (1886:173-175)  states 
that  he  has  often  found  larvae  in  the  wet  detritus  underneath  the  bark 


988  CHARLES  PAUL  ALEXANDER 

of  decaying  trees.  The  larvae  are  much  closer  to  Ctenophora  than  to 
Tanyptera,  the  skin  being  tough  and  opaque,  with  a  fine  pubescence,  and 
the  spiracular  disk  consisting  of  small  but  distinct  lobes.  The  pupae  like- 
wise are  similar  to  those  of  Ctenophora,  having  the  pronotal  breathing 
horns  elongate,  five  spines  on  abdominal  sternites  3  to  6,  and  four  spines  on 
tergites  2  to  7. 

Genus  Tanyptera  Latreille  (Gr.  extend  4-  wing) 

1805     Tanyptera  Latr.     Hist.  Nat.  Crust,  et  Ins.,  vol.  14,  p.  286. 
1832     Xiphura  Brulle".     Ann.  Soc.  Ent.  France,  vol.  1,  p.  206. 

Larva. —  Integument  very  thin,  with  numerous  setae,  those  on  dorsum  very  small. 
Spiracular  disk  with  lobes  practically  lacking.  Spiracles  large,  lying  exposed  on  the  face  of 
last  segment.  Anal  gills  bluntly  rounded.  Mandible  small,  with  one  dorsal  and  one  ventral 
tooth.  Antenna  cylindrical,  capped  with  an  apical  cone.  Mentum  with  seven  to  nine 
teeth. 

Pupa. —  Cephalic  crest  lacking.  Sheaths  of  maxillary  palpi  not  recurved  at  tips.  Pronotal 
breathing  horns  large,  broadly  flattened,  margin  deeply  crenulated.  Mesonotum  with  two 
blunt  tubercles.  Abdominal  segments  with  six  to  ten  spines  on  tergites,  three  to  five  spines 
on  sternites. 

Tanyptera  is  a  small  genus  including  about  twelve  nominal  species 
whose  limits  and  relationships  are  still  but  little  understood.  The  imma- 
ture stages  are  spent  in  the  decayed  or  partly  decayed  wood  of  various 
deciduous  trees. 

In  Europe,  Tanyptera  atrata  (Linn.),  the  genotype,  was  found  by  Ferris 
(1840)  and  by  De  Rossi  (1882)  in  decaying  alder  (Alnus)  stems.  Nord- 
linger  (1848)  found  the  same  species  in  linden  (Tilia)  and  in  poplar 
(Populus).  It  has  also  been  taken  in  oak  (Quercus),  beech  (Fagus), 
birch  (Betula),  and  other  hardwood  species.  Gerbig  (1913)  discussed  the 
variety  ruficornis  Meig.  under  the  name  Ctenophora  flavicornis. 

In  America,  Malloch  (1915-17  b:  194-195)  describes  T.  fumipennis 
(O.  S.)  from  a  much-decayed  chestnut  log  (Castanea),  and  later  (1919) 
in  basswood  (Tilia),  where  the  species  was  associated  with  larvae  of 
Xylota  fraudulosa  Loew  and  Chalcomyia  aerea  (Loew),  of  the  family 
Syrphidae.  Tanyptera  frontalis,  discussed  below,  was  found  in  red  maple 
(Acer). 

Tanyptera  frontalis  (0.  S.) 

1864    Ctenophora  frontalis  O.  S.     Proc.  Ent.  Soc.  Phila,  vol.  3,  p.  48. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  989 

The  writer  found  numerous  larvae  of  Tanyptera  frontalis  in  a  fallen 
log  of  red  maple  (Acer  rubrum  Linn.)  near  Beebe  Lake,  Ithaca,  New 
York,  on  March  22,  1913.  Larvae  of  several  sizes  were  found.  They 
were  working  in  wood  which  was  well  preserved,  not  entirely  sound  but 
still  so  hard  that  it  had  to  be  cut  with  a  hatchet.  The  larvae  pupated 
in  April.  A  small  male  emerged  on  May  1. 

Other  larvae  of  Tanyptera  were  found  in  a  hickory  log  (Carya  sp.) 
at  Sandy  Landing,  Virginia  (opposite  Plummers  Island),  on  September 
9,  1913.  Detailed  drawings  of  the  larval  structure  made  by  Dr.  Boving 
are  in  the  collection  of  the  United  States  National  Museum. 

Larva. —  Length,  30-35  mm. 
Diameter,  7-7.2  mm. 

Coloration,  pale  yellowish  white. 

Form  terete,  very  stout.  Integument  thin.  Numerous  long  black  setae  on  segments 
(Plate  LXXXVIII,  482),  arranged  in  transverse  rows  before  posterior  margin;  setae  of 
dorsum  (Plate  LXXXVIII,  481)  very  tiny,  one  on  either  side  of  median  line;  a  pair  of  setae 
laterad  of  these  and  in  alinement;  setae  on  pleural  region  very  long  and  delicate;  on  thoracic 
segments,  setae  at  about  midlength;  on  abdominal  segments,  setae  closer  to  posterior 
margin;  mid- ventral  setae  very  tiny,  four  in  number,  on  thoracic  segments  at  about  mid- 
length,  forming  a  stiff  pencil  on  sides  of  venter,  with  two  small  setae  between;  laterad  of 
these  four  intermediate  setae,  a  group  of  three  setae,  two  long  and  one  very  short;  ventral  setae 
lying  at  a  level  posterior  to  that  of  pleural  setae.  Spiracular  disk  (Plate  LXXXVIII, 
483)  with  lobes  practically  lacking,  the  two  large,  oval  spiracles  lying  exposed  on  trun- 
cated end  of  last  segment;  above  and  laterad  of  each  spiracle,  a  small,  blunt  lobe  with  a 
blackened  mark  and  three  long  setae;  below  spiracles,  two  narrow  black  lines  representing 
the  two  ventral  lobes;  a  small  pencil  of  setae  below  each  of  these  marks,  and  each  mark 
having  a  sensory  bristle;  three  or  four  long  setae  on  sides  of  spiracular  disk.  Spiracles  with 
small  middle  piece  black,  ring  yellowish  brown;  spiracles  separated  by  a  distance  a  little 
greater  than  diameter  of  one.  Anal  gills  four,  bluntly  rounded,  very  protuberant  ,and 
evidently  formed  for  propulsion.  (In  older  specimens  the  anal  gills  are  minutely  roughened 
and  are  darker  in  color.) 

Head  capsule  of  the  massive  tipuline  type.  Labrum  rather  broad;  median  epipharyngeal 
region  with  a  small  brush  of  hairs  surrounding  two  sensory  setae;  lateral  lobes  large,  on  their 
ventral  face  densely  hairy,  surrounding  four  sensory  setae  and  a  flattened  hyaline  peg. 
Mentum  (Plate  LXXXVIII,  478)  rather  small,  broadly  rounded,  anterior  margin  with 
seven  or  nine  teeth,  in  the  latter  case  the  outermost  tooth  on  either  side  very  blunt  and 
reduced,  the  median  tooth  long  and  flattened.  Antenna  (Plate  LXXXVIII,  479)  short- 
cylindrical;  apical  papilla  very  small,  hyaline,  conical,  with  surface  sculptured;  in  addition 
to  this  cone,  three  or  four  small  hyaline  sense  pegs;  the  usual  auditory  organ  located  at 
about  midlength  of  segment.  Mandible  (Plate  LXXXVIII,  480)  small,  with  one  dorsal  and 
one  ventral  tooth  in  addition  to  the  apical  point,  ventral  tooth  flattened  and  with  margin 
crenulated;  a  stout  seta  at  heel  of  mandible;  prostheca  an  elongated  cone,  situated  at  base 


990  CHARLES  PAUL  ALEXANDER 

of  mandible.  Maxilla  rather  small,  simple;  palpus  large,  antenniform,  with  apex  bluntly 
rounded;  inner  lobe  densely  covered  with  short,  stout  setae  surrounding  a  powerful  bristle 
and  a  small  brown  sensory  organ. 

Pupa—  Length,  30-33  mm. 

Width,  d.-s.,  5-6  mm. 
Depth,  d.-v.,  5.5  mm. 

Coloration  pale  yellow;  pronotal  breathing  horns  liver-colored.  (In  older  specimens, 
the  thorax  and  appendages  are  dark-colored,  and  the  abdomen  has  broad  brown  sublateral 
stripes.) 

Head  rather  small,  cephalic  crest  lacking.  Antenna  stout,  rather  elongate,  extending  far 
beyond  ends  o '  palpi,  segments  angulated.  Clypeus  and  labrum  tumid,  transversely  wrinkled. 
Labial  lobes  slender,  divergent.  Sheaths  of  maxillary  palpi  rather  slender,  tips  curved 
but  not  recurved  (Plate  LXXXVIII,  485).  Pronotal.  breathing  horn  (Plate  LXXXVIII, 

486)  large,  broadly  flattened,  slightly  incurved,  broader  at  apex  than  at  base,  deeply  furrowed 
up  middle  of  outer  face,  the  broad  margin  thus  formed  deeply   wrinkled  to  crenulate. 
Mesonotum  large,  transversely  wrinkled,  with  two  blunt  tubercles  provided  with  short  setae 
(Plate  LXXXVIII,  484).     Wing  sheaths  small,  reaching  end  of  second  abdominal  segment. 
Leg  sheaths  ending  before  tip   of  third  abdominal  segment;  fore  tarsi  very  short,   hind 
tarsi  the  longest,  those  of  middle  legs  intermediate. 

First  abdominal  tergite  with  two  spines;  segments  2  to  7  broad,  divided  into  the  usual  basal 
and  posterior  rings;  second  segment  on  posterior  ring  with  four  spines;  segments  3  to  6  with 
six  to  ten  spines,  the  intermediate  ones  usually  smaller;  segment  7  with  four  spines;  pleurites 
with  a  spine  on  each  ring;  sternites  with  similar  arrangement  to  that  of  tergites,  but  spines 
usually  fewer  in  number,  segments  3  and  4  with  only  a  single  widely  separated  spine  on  each 
side,  segments  5  and  6  with  four  or  five  spines,  segment  7  with  three  spines;  segment  8  has 
four  small  spines  between  the  large  lateral  ones  described  below.  Male  cauda  blunt,  dorsal 
lobes  very  divergent,  ending  in  sharp,  chitinized  points.  Female  cauda  (Plate  LXXXVIII, 

487)  with  tergal  valves  elongate,  narrowed  to  the  moderately  acute  tips;  sternal  valves 
similar  in  shape  but  smaller;  lateral  lobes  of  ninth  segment  directed  caudad  and  laterad, 
at  tips  running  out  into  chitinized  points;  two  small  setae  before  tips;  segment  8  with  a 
powerful  lateral  lobe  on  either  side,  each  terminating  in  a  chitinized  point;  posterior  lat- 
eral angles  of  segment  produced  into  slender,  blunt  points. 

Nepionotype.—  Ithaca,  New  York,  March  22,  1913. 
Neanotype.— May  1,  1913.     No.  11-1913. 
Paratypes. —  Four  larvae  and  two  pupae  with  types. 

Subtribe  Tipularia 

Genus  Longurio  Loew  (Lat.  a  tall  man) 

1869    Longurio  Loew.    Berl.  Ent.  Ztschr.,  vol.  13,  p.  2. 

Larva  (supposition).' — Body  massive.  Integument  semi-transparent.  Form  clearly 
depressed.  Spiracular  disk  surrounded  by  six  lobes,  dorsal  pair  very  small,  ventral  pair 
very  long;  ventral  and  lateral  lobes  provided  with  but  few  setae  at  and  near  tips;  spiracular 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  991 

disk  and  lobes  unmarked.  Spiracles  small.  Anal  gills  branched.  Mandible  small,  with 
but  a  single  dorsal  and  ventral  tooth  in  addition  to  apical  point.  Mentum  with  nine  teeth. 
Hypopharynx  five-toothed. 

Pupa.- —  Antenna  short,  ending  opposite  tips  of  maxillary  palpi.  Sheaths  of  maxillary 
palpi  not  recurved  at  tips.  Mesonotum  unarmed.  Pronotal  breathing  horns  very  long  and 
slender,  one,  at  least,  about  half  length  of  body.  Wings  showing  clearly  the  characteristic 
venation  of  Longurio.  Abdomen  elongate,  posterior  ring  of  individual  segments  with  short, 
stout  spines,  including  a  few  on  pleurites;  dorsum  of  eighth  segment  with  four  powerful  lobes. 

Longurio  is  a  small  genus  including  about  ten  described  species,  of 
which  two  occur  in  eastern  North  America.  The  genotype,  Longurio 
testaceus,  is  the  best-known  species  locally.  The  immature  stages  of  this 
species  are  here  discussed  for  the  first  time.  They  are  spent  in  sand 
or  sandy  earth.  The  branched  anal  gills  of  the  larva,  and  the  excessively 
elongate  breathing  horns  of  the  pupa,  are  notable  features. 

Longurio  testaceus  Loew 

1869    Longurio  testaceus  Loew.    Berl.  Ent.  Ztschr.,  vol.  13,  p.  2. 

Longurio  testaceus  is  probably  the  largest  crane-fly  in  North  America, 
the  females  exceeding  the  better-known  Holorusia  of  the  Western  States. 
The  adult  flies  are  difficult  to  capture,  being  very  wary.  When  the  insect 
is  at  rest  the  body  generally  hangs  perpendicularly,  with  the  wings  folded 
incumbent  over  the  abdomen. 

On  November  9,  1916,  Mr.  Hyslop  sent  the  writer  two  living  larvae 
which  are  referred  with  little  doubt  to  this  species.  They  were,  found 
in  wet  sand  in  a  bog  on  the  top  of  South  Mountains,  near  Myersville, 
Maryland.  The  larvae  were  very  restless,  the  head  capsule  being  con- 
stantly exserted  and  withdrawn.  Waves  of  contraction  start  from  the 
posterior  end  of  the  body  and  pass  toward  the  head.  A  large  pupa  taken 
by  Dr.  J.  C.  Bradley  at  Tallulah  Falls,  Georgia,  on  June  17,  1910, 
undoubtedly  belongs  to  this  species,  the  venation  being  clearly  apparent 
on  the  wing  pad.  An  additional  cast  pupal  skin  is  in  the  collection 
of  the  United  States  National  Museum. 

Larva  (supposition).— Length,  31  mm.  contracted,  58  mm.  extended. 
Diameter,  11-12.5  mm. 

Color  whitish,  subhyaline;  thoracic  segments  more  yellowish;  in  life  the  brown  food  contents 
showing  clearly  thru  abdomen. 

Form  very  depressed,  lateral  folds  prominent;  body  very  stout  and  fleshy.     Skin  very  tl 
and  semitransparent,  showing  internal  organs  within,  practically  destitute  of  pubescence; 


992  CHARLES  PAUL  ALEXANDER 

a  few  short  setae  on  body,  especially  on  thoracic  segments.  Spiracular  disk  (Plate  LXXXIX, 
490)  large,  flattened,  surrounded  by  six  lobes;  dorsal  pair  very  small,  represented  by  two  short, 
conical  protuberances;  lateral  and  ventral  lobes  slender,  the  latter  a  little  the  longer,  each 
with  three  or  four  long,  delicate  setae  at  tip  and  two  or  three  others  before  tip  on  outer  face; 
a  long  seta  on  margin  of  disk  between  dorsal  and  lateral  lobes;  spiracular  disk  and  lobes 
entirely  unmarked  with  darker.  Spiracles  small,  circular,  stigmal  rings  very  narrow;  spir- 
acles separated  by  a  distance  equal  to  about  three  times  diameter  of  one.  Anal  gills  four, 
large,  pinnately  branched,  each  gill  with  six  lateral  branches. 

Head  capsule  massive,  of  the  tipuline  type.  Labrum  with  a  distinct,  densely  hairy  lobe 
on  either  side.  Mentum  (Plate  LXXXIX,  488)  large;  anterior  margin  with  nine  slender 
teeth,  median  one  the  longest,  outermost  teeth  on  either  side  blunt,  flattened,  evidently 
formed  by  fusion  of  two  teeth.  Hypopharynx  (Plate  LXXXIX,  489)  five-toothed;  teeth 
blunt,  the  three  middle  ones  larger,  the  lateral  tooth  on  either  side  a  little  smaller.  Antenna 
long  and  slender,  cylindrical;  apex  blunt  and  without  distinct  sense  pegs  or  setae.  Mandible 
small,  with  a  large  conical  dorsal  tooth  and  a  single  flattened  ventral  tooth  in  addition 
to  apical  point;  prostheca  distinct.  Maxilla  small;  palpi  large,  cylindrical,  truncated  at 
apex  and  with  a  circular  auditory  plate  near  end;  inner  lobe  of  maxilla  with  abundant  elon- 
gate setae. 

Pupa. —  Length,  42-45  mm. 

Length  of  longest  breathing  horn,  18-19  mm.  additional. 
Width,  d.-s.,  3  mm. 
Depth,  d.-v.,  4.2  mm. 

Color  dark  brown;  pronotal  breathing  horns  paler  at  tips;  mesonotum  chestnut  brown; 
abdominal  segments  indistinctly  ringed  with  pale  and  darker. 

Front  between  antennal  bases  swollen,  finely  tuberculate  but  without  distinct  setae. 
Antenna  very  short,  ending  opposite  tips  of  maxillary  palpi.  Labrum  large.  Sheaths  of 
maxillary  palpi  not  recurved  at  tips.  Mesonotum  unarmed,  with  fine  transverse  wrinkles. 
Pronotal  breathing  horns  (Plate  LXXXIX,  491)  very  long  and  slender,  the  right  one,  at  least, 
exceedingly  elongate,  with  tip  expanded.  (The  left  breathing  horn  was  broken  before  the 
apex  in  both  the  pupae  studied;  it  was  almost  as  long  as  the  right  horn,  and  may,  of  course, 
have  been  longer.)  Wing  pads  reaching  end  of  second  abdominal  segment;  characteristic 
venation  of  genus  showing  clearly  on  sheath.  Leg  sheaths  long,  extending  to  beyond  mid- 
length  of  fourth  abdominal  segment;  fore  legs  shorter  than  the  others. 

Abdomen  elongate.  Abdominal  segments  2  to  7  near  posterior  margin  with  a  transverse 
row  of  short,  stout  spines  which  are  interrupted  only  near  pleura;  pleural  area  with  four 
or  five  spines;  ventral  and  dorsal  segments  with  numerous  spines;  dorsal  row  of  spines  more 
distant  from  posterior  margin  of  segment  than  the  other  areas;  dorsum  of  eighth  segment 
with  four  powerful,  chitinized  lobes  directed  dorsad  and  caudad;  posterior  margin  of  these 
lobes  with  about  five  or  six  small  teeth;  two  spines  near  base  of  pleural  region  on  segment  8. 
Male  cauda  consisting  of  two  blunt  sheaths,  lying  between  posterior  pair  of  lobes  described 
above. 

Larva. —  South  Mountains,  near  Myersville,  Maryland,  November  6,  1916. 
Neanotype. —  Tallulah  Falls,  Georgia,  June  17,  1910. 
Paratype. —  Cabin  John  Bridge,  Maryland,  May  31,  1900. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  993 

Genus  Aeshnasoma  Johnson  (Gr.  a  dragon  fly  +  body) 

1909    Aeshnasoma  Johns.    Proc.  Boat.  Soc.  Nat.  Hist.,  vol.  34,  p.  115-116. 

Aeshnasoma  is  a  monotypic  genus  found  in  northeastern  North  America. 
It  is  unquestionably  close  to  Longurio  and  may  be  congenerous  with  it. 
The  type,  Aeshnasoma  rivertonensis  Johns.,  is  apparently  very  local  in 
its  distribution. 

Johnson  (1906 : 1-2)  described  an  unknown  tipuline  larva  which  undoubt- 
edly pertains  to  this  species.  The  larva  was  found  on  June  10,  1900, 
in  a  cold  spring  at  Riverton,  New  Jersey.  It  was  brought  into  the  labora- 
tory but  could  not  be  reared,  the  change  from  the  cold  spring  (about 
60°  F.)  to  warmer  waters  being  fatal.  The  larva  when  fully  extended 
measured  about  45  millimeters  in  length.  It  was  yellowish  white  in 
color  and  was  translucent,  the  alimentary  canal  with  its  contents  being 
clearly  visible  thru  the  thin  skin.  Johnson  describes  and  figures  the 
peculiar  branched  anal  gills  (Plate  LXXXIX,  492)  of  this  genus.  The 
larva  was  doubtfully  referred  to  Longurio,  the  adults  of  Aeshnasoma 
being  undescribed  at  that  time. 

In  a  later  paper  (1907-12  [1909] :  115-116)  Johnson  mentions  the  taking 
of  several  more  larvae  in  1902,  and,  on  July  20,  the  capture  of  the  adult  flies 
on  which  the  genus  and  species  are  based. 

The  only  larva  that  was  preserved  was  kindly  sent  to  the  writer  for 
study  by  Mr.  Johnson.  It  is  undoubtedly  very  close  to  Longurio,  both 
genera  showing  the  same  peculiar  spiracular  disk  and  the  branched  anal 
gills,  a  condition  that  is  found  nowhere  else  in  the  Tipulidae  so  far  as  is 
known  to  the  writer. 

Genus  Holorusia  Loew  (derivation  obscure) 

1863    Holorusia  Loew.    Berl.  Ent.  Ztschr.,  vol.  7,  p.  1. 

Larva. —  Spiracular  disk  surrounded  by  six  moderately  elongate  lobes  fringed  with  long 
hairs;  inner  face  of  lateral  and  ventral  lobes  with  capillary  black  lines;  disk  between  spiracles 
dusky.  Anal  gills  six.  Mandible  small,  with  a  single  dorsal  and  ventral  tooth  in  addition 
to  apical  point.  Antenna  with  a  conical  apical  papilla.  Mentum  seven-toothed.  Hypo- 
pharynx  six-toothed. 

Pupa.  —  Sheaths  of  maxillary  palpi  recurved  at  tips.  Pronotal  breathing  horns  rather 
short  and  stout,  the  long  apices  flattened.  Armature  of  abdominal  segments  almost  as  in 
Prionocera,  but  the  posterior  rows  of  spines  more  numerous  (fourteen  to  twenty-four); 
pleurites  with  three  spines,  the  two  on  posterior  ring  situated  one  behind  the  other.  Cauda 
with  six  stout  dorsal  lobes. 


994  CHARLES  PAUL  ALEXANDER 

Holorusia  is  a  small  genus  (about  ten  species)  of  New  World  crane- 
flies,  only  one  of  which — -the  genotype,  Holorusia  rubiginosa  —  is  Nearctic. 
This  species  and  Longurio  are  the  largest  Nearctic  crane-flies.  The 
anatomy  of  the  " giant  crane-fly"  has  been  described  in  some  detail 
by  Kellogg  (1901,  a  and  b)  and  by  Comstock  and  Kellogg  (1904).  The 
immature  stages  are  spent  in  moist  earth.  The  genus  is  undoubtedly 
closely  related  to  Prionocera,  and,  presumably,  to  the  Old  World  genus 
Ctenacroscelis  Enderlein. 

Holorusia  rubiginosa  Loew 

1863     Holorusia  rubiginosa  Loew.     Berl.  Ent.  Ztschr.,  vol.  7,  p.  1. 
1888     Tipula  (Holorusia)  grandis  Bergr.     Ent.  Tidskr.,  vol.  9,  p.'  140. 

Holorusia  rubiginosa  is  widely  distributed  thruout  the  western  United 
States  and  Canada.  A  number  of  larvae  were  taken  by  H.  Morrison 
near  Stanford  University,  California,  on  February  22,  1915.  They  were 
shipped  to  the  writer  at  Ithaca,  New  York,  where  the  species  was  reared. 
The  massive  larva  is  used  for  purposes  of  dissection  in  the  entomological 
courses  at  some  of  the  western  universities. 

Larva. —  Length,  50-60  mm. 

Diameter,  6.2-6.4  mm. 

Coloration,  dark  greenish  brown. 

Form  stout,  subterete.  Integument  covered  with  dense,  short,  erect,  black  hairs.  A 
few  weak  and  delicate  setae,  two  on  dorsum  and  on  venter  of  each  abdominal  segment;  two 
long  setae  on  lateral  margins  of  posterior  rings.  Spiracular  disk  (Plate  XC,  496)  moderately 
large,  surrounded  by  six  stout,  elongate  lobes  which  are  similar  to  those  of  Prionocera  but 
are  stouter  and  less  digitiform;  ventral  lobes  a  little  the  longest,  dorsal  lobes  a  little  the  short- 
est; all  the  lobes  capable  of  close  approximation,  completely  protecting  spiracles;  lobes 
fringed  with  long  black  hairs  which  are  longest  near  apices,  shorter  between  lobes;  ventral 
and  lateral  lobes  with  a  delicate  black  line  down  inner  face,  these  lines  barely  indicated 
on  dorsal  lobes;  remainder  of  disk  and  lobes  dusky.  Spiracles  very  large,  circular,  separated 
by  a  distance  a  little  less  than  diameter  of  one.  Anal  gills  six,  short,  slender,  the  two 
anterior  gills  of  either  side  united  basally,  posterior  pair  simple. 

Head  capsule  of  the  usual  massive  tipuline  type,  prefrons  running  caudad  as  a  narrow 
point,  lateral  plates  broad.  Labrum  broad,  with  a  densely  hairy  lobe  on  either  side.  Men- 
turn  (Plate  XC,  493)  with  a  prominent  median  point;  behind  it  on  either  side  three  flattened 
teeth,  the  innermost  the  broadest,  the  middle  tooth  more  acute,  the  outermost  formed  by 
fusion  of  two  small  teeth.  Hypopharynx  (Plate  XC,  494)  about  six-toothed,  the  intermediate 
teeth  with  a  large  notch  between.  Antenna  (Plate  XC,  495)  with  basal  segment  very  long 
and  slender;  principal  apical  papilla  conical;  a  number  of  small  hyaline  sense  pegs.  Mandi- 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  995 

ble  slender,  with  only  two  teeth,  a  stout  dorsal  tooth  and  a  single  flattened  ventral  tooth. 
Maxilla  small,  lobes  covered  with  short,  dense  hairs. 

Pupa. —  Length,  32  mm. 

Width,  d.-s.,  5  mm. 
Depth,  d.-v.,  5  mm. 

Color  brown;  flattened  lateral  margins  of  abdomen  broadly  yellowish. 

Thorax  terete;  abdomen  depressed,  with  lateral  margins  flattened,  carinate.  Cephalic 
crest  represented  by  two  low,  parallel  ridges,  provided  with  one  or  two  tiny  setae.  Lab  rum 
large,  tumid,  transversely  wrinkled,  the  blunt  apex  completely  separating  the  diamond- 
shaped  labial  lobes.  Maxillary  palpi  stout,  extreme  tip  recurved.  Antenna  moderately 
elongated,  extending  some  distance  beyond  maxillary  palpi.  Pronotal  breathing  horns  rather 
short  and  stout,  finely  ringed,  the  rather  long  tips  flattened,  about  equal  to  one-fifth  length 
of  entire  organ.  Mesonotum  convex  (Plate  XC,  497),  with  transverse  anastomosing 
wrinkles;  on  either  side  of  median  line  behind,  a  blunt  tubercle.  Wing  sheaths  reaching  end 
of  second  abdominal  segment.  Leg  sheaths  extending  just  beyond  base  of  fourth  abdom- 
inal segment;  fore  tarsi  short,  middle  tarsi  a  little  longer  than  hind  tarsi. 

Abdominal  segments  with  the  usual  basal  and  posterior  rings;  armature  almost  as  in 
Prionocera;  tergites  with  posterior  row  of  spines  numbering  between  twenty  and  twenty- 
four  on  intermediate  segments,  near  anterior  lateral  angle  two  spines,  basal  ring  unarmed; 
pleurites  with  one  setiferous  spine  on  basal  ring  and  two  similar  spines  on  posterior  ring, 
one  placed  considerably  behind  the  other;  sternites  armed  similarly  to  tergites,  but  the  poste- 
rior row  of  spines  larger  and  somewhat  fewer  in  number  (fourteen  to  eighteen) ;  posterior  ring 
on  either  side  median  line  near  base  with  two  spines,  the  innermost  very  large  and  power- 
ful. Male  cauda  with  ventral  lobes  blunt,  each  armed  with  a  slender  black  spine  near  pos- 
terior margin;  dorsal  surface  of  cauda  almost  as  in  Prionocera,  armed  with  six  stout  lobes, 
which  here  are  shorter  and  stouter,  with  tips  abruptly  narrowed;  lateral  margin  of  segment 
8  with  a  stout  lobe  on  either  side,  each  terminating  in  a  cylindrical  spine.  Female  cauda 
similar  to  male  cauda,  dorsum  with  the  same  six  lobes;  acidothecae  short,  tergal  valves 
slightly  exceeding  the  more  blunt  sternal  valves.  ; 

Nepionotype. —  Stanford  University,  California,  February  27,  1915. 

Neanotype. —  April  5,  1915. 

Paratypes. —  Larvae  and  pupae  with  types. 

Genus  Prionocera  Loew  (Gr.  saw  +  horn) 

1844     Prionocera  Loew.     Stett.  Ent.  Zeit.,  vol.  5,  p.  170. 
1863    Stygeropis  Loew.     Berl.  Ent.  Ztschr.,  vol.  7,  p.  298. 

Larva.—  Spiracular  disk  surrounded  by  six  long,  finger-like  lobes  fringed  with  long,  deli- 
cate hairs;  each  lobe  with  a  capillary  black  line  down  middle  of  inner  face.  Spiracles  large. 
Anal  gills  unbranched.  Mentum  seven-  to  nine-toothed.  Hypopharynx  five-toothed. 
Mandible  with  about  two  dorsal  and  three  ventral  teeth. 

Pupa.— Maxillary  palpi  recurved  at  tips.  Pronotal  breathing  horns  very  elongated, 
unequal,  the  longer  one  about  half  length  of  body;  horns  at  tips  split  into  long  flaps.  Abdom- 
inal tergites  with  a  posterior  transverse  row  of  fifteen  or  fewer  spines,  and  two  small  spines 


996  CHARLES  PAUL  ALEXANDER 

near  anterior  lateral  margin  of  posterior  ring;  pleurites  with  a  spine  on  basal  ring  and  two 
transverse  spines  on  posterior  ring.     Cauda  with  six  strong  dorsal  lobes. 

Prionocera  is  a  small  genus  (about  a  dozen  species)  of  usually  far  northern 
flies  of  somber  coloration.  The  only  species  found  in  eastern  North 
America  is  Prionocera  fuscipennis,  discussed  below.  The  immature 
stages  are  somewhat  similar  to  those  of  Holorusia.  The  apparent  similarity 
of  the  pupa  to  that  of  Longurio  is  probably  not  indicative  of  a  very  close 
relationship. 

The  immature  stages  have  been  discussed  but  little  in  the  literature. 
The  "Tipula  sp.  No.  1"  of  Malloch  (1915-17  b:  199-200)  refers  to  P.  fusci- 
pennis.  The  immature  stages  of  a  species  supposed  to  be  P.  parri  (Kirby) 
have  been  discussed  and  figured  by  the  writer  in  his  report  on  the  Canadian- 
Arctic  Tipulidae  (Alexander,  1919  c:  19c-20c). 

The  name  Stygeropis  has  been  in  use  for  many  years  under  the  belief 
that  the  earlier  name  Prionocera  was  preoccupied  in  the  Coleoptera. 
Dr.  Bergroth  states  that  this  is  not  so  and  that  Prionocera  should  be 
used. 

Prionocera  fuscipennis  (Loew) 

1865    Stygeropis  fuscipennis  Loew.    Berl.  Ent.  Ztschr.,  vol.  9,  p.  129. 

C.  H.  Kennedy  found  two  cast  pupal  skins  among  Sparganium  stems 
in  Ringwood  Hollow,  Ithaca,  New  York,  on  November  20,  1916.  Several 
larvae  had  been  found  here  in  the  preceding  July,  and  some  others  were 
found  on  June  4,  1917  (No.  106-1917),  in  a  cat-tail  swamp  near  Bool's 
hillside,  Ithaca,  where  they  were  associated  with  the  characteristic  helo- 
phytic  crane-fly  fauna  (Bittacomorpha,  Rhamphidia  flavipes,  Pseudolim- 
nophila  luteipennis,  Pilaria  recondita,  Tipula  tricolor,  and  other  species). 
Malloch's  material  was  taken  in  Wisconsin  in  May.  Dr.  Needharn  has 
reared  the  species  near  Lake  Forest,  Illinois. 

Larva. —  Length,  18-22  mm. 
Diameter,  2-2.2  mm. 

Coloration  dark  brown,  in  some  cases  with  a  pale  dorse-median  stripe. 

Form  terete,  tapering  gradually  to  anterior  end  of  body.  Segments  with  several  scattered 
elongate  setae.  Spiracular  disk  (Plate  XCI,  502)  surrounded  by  six  long,  finger-like  lobes 
which  are  delicately  fringed  with  long  hairs;  ventral  lobes  considerably  the  longest;  lateral 
lobes  a  little  larger  and  stouter  than  dorsal  lobes;  all  the  lobes  broadly  margined  with  dark 
brown,  these  marks  expanding  at  inner  ends;  on  ventral  lobes,  lateral  margin  expanded  at  inner 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  997 

end  and  continued  across  disk,  meeting  its  fellow  of  the  opposite  side  between  spiracles; 
dorsal  margin  of  lateral  lobes  touching  spiracles;  each  of  the  lobes  marked  with  a  capillary 
dark  brown  line  down  center  of  inner  face,  this  beginning  near  ends  of  lobes  and  extending 
almost  to  base;  lobes  fringed  with  long  hairs,  these  very  tiny  near  base,  longer  near  tips  of 
lobes,  but  scarcely,  if  at  all,  interrupted  between  lobes.  Spiracles  large,  circular,  sepa- 
rated by  a  distance  a  little  greater  than  diameter  of  one.  Anal  gills  six,  long,  slender, 
unbranched. 

Head  capsule  of  the  usual  tipuline  type.  Labrum  covered  with  dense,  short  hairs,  those 
on  lateral  lobes  longer.  Mentum  (Plate  XCI,  498)  usually  seven-toothed,  in  some  cases  nine- 
toothed;  median  tooth  slender,  lateral  teeth  flattened,  subacute.  Hypopharynx  (Plate 
XCI,  499)  narrow,  five-toothed.  Antenna  (Plate  XCI,  500)  long  and  slender;  first  segment 
a  little  enlarged  near  base;  at  apex  several  tiny  sensory  papillae;  auditory  plate  near  base 
of  segment.  Mandible  (Plate  XCI,  501)  moderately  large,  with  two  large  dorsal  and  about 
three  ventral  teeth;  prostheca  large.  Maxilla  small;  palpus  large,  cylindrical,  apex  truncated. 

Pupa. —  Length,  15-22  mm. 
Width,  d.-s.,  2.4  mm. 
Depth,  d.-v.,  2.6  mm. 

Coloration  dark  brown;  lateral  and  posterior  margins  of  abdominal  segments  paler.  (In 
old  specimens  the  general  coloration  is  very  dark  brown;  in  younger  specimens  the  abdomen 
is  more  or  less  distinctly  lined  with  brown.) 

Thorax  subterete;  abdomen  depressed,  lateral  margins  flattened.  Labrum  broad,  apical 
point  narrow.  Labial  lobes  broad,  slightly  separated  on  median  line.  Maxillary  palpi 
short,  stout,  apex  recurved.  Antenna  moderately  elongated,  extreme  tip  darkened.  Pro- 
notal  breathing  horns  long  and  slender,  unequal  in  length,  the  longer  about  9  or  10  mm. 
in  length,  the  other  6  mm.,  at  tips  split  into  divergent  flaps  (Plate  XCI,  504)  almost  as  in 
the  hexatomine  genera  Pseudolimnophila  and  Pilaria,  which  live  in  the  same  muddy  sit- 
uations. Mesonotum  (Plate  XCI,  503)  transversely  wrinkled.  Leg  sheaths  reaching  posterior 
margin  of  third  abdominal  segment;  hind  legs  the  longest;  middle  legs  a  little  shorter  than 
fore  legs. 

Abdominal  segments  divided  into  a  basal  and  a  posterior  ring;  tergites  with  basal  ring 
unarmed;  posterior  ring  with  a  subterminal  transverse  row  of  short  spines,  with  a  few  setae 
located  on  lateral  face  of  some  of  the  spines;  on  second  tergite,  four  to  six  spines,  on  ter- 
gites 3  to  7,  three  to  fifteen  spines;  two  small  spines  with  setae  near  anterior  lateral  angle 
of  posterior  ring;  pleurites  with  a  small  setiferous  spine  on  basal  ring,  and  two  such  spines 
on  posterior  ring  located  side  by  side;  sternites  with  the  basal  ring  unarmed,  posterior  ring 
armed  similarly  to  that  of  tergites;  in  addition  to  posterior  row  of  spines,  a  pale  oval  area 
on  either  side  of  midventral  line,  each  with  two  transversely  placed  spines.  Male  cauda 
(Plate  XCI,  505)  with  four  powerful  lobes  on  dorsum  of  last  segment,  directed  dorsad  and 
slightly  caudad,  lobes  bearing  three  or  four  small  spines  before  tips;  between  anterior  pair 
of  lobes,  two  additional  slender  lobes,  each  ending  in  two  acute  spines. 

Nepionotype. —  Ringwood  Hollow,  Ithaca,  New  York,  July  20,  1916. 
Neanotype. —  Cast  pupal  skin,  type  locality,  November  20,  1916. 
Paratypes. —  Larvae  and  pupal  skins,  type  locality. 


998  CHARLES  PAUL  ALEXANDER 

Genus  Tipula  Linnaeus  (Lat.  a  water-strider) 

1758  Tipula  Linn.    Syst.  Natur.,  ed.  10,  p.  585. 

1842  Pterelachisus  Rond.     Mag.  Zool.  Ins.,  pi.  106. 

1864  Anomaloptera  Lioy.     Atti  dell'  Institut  Veneto,  ser.  3,  vol.  9,  p.  218. 

1887  Oreomyza  Pokorny.     Wien.  Ent.  Ztg.,  vol.  6,  p.  50. 

1894  Manapsis  Scudder.     Proc.  Amer.  Philos.  Soc.,  vol.  32,  p.  222. 

1894  Rhadinobrochus  Scudder.     Proc..  Amer.  Philos.  Soc.,  vol.  32,  p.  223. 

1894  Tipulidea  Scudder.     Proc.  Amer.  Philos.  Soc.,  vol.  32,  p.  238-239. 

1916  Nippotipula  Mats.    Thous.  Ins.  Japan,  add.  2,  p.  457-458. 

1916  Platytipula  Mats.     Thous.  Ins,  Japan,  add.  2,  p.  459. 

1916  Yamatotipula  Mats.     Thous.  Ins.  Japan,  add.  2,  p.  461-462. 

1916  Togotipula  Mats.     Thous.  Ins.  Japan,  add.  2,  p.  465. 

Larva.- —  Form  generally  stout,  terete  or  nearly  so.  Integument  with  pubescence  and 
almost  invariably  with  a  definite  chaetotaxy.  Spiracular  disk  surrounded  by  six  or  rarely 
eight  lobes,  simple,  or  in  certain  species  (as  T.  abdominalis)  more  or  less  split  at  their  tips. 
Spiracles  small  and  widely  separated  (in  T.  abdominalis},  or  in  other  species  large  and  rather 
close  together.  Anal  gills  almost  invariably  present,  with  six  or  eight  branches,  these 
branches  simple,  not  pinnate.  Head  capsule  compact  and  massive.  Labrum  broadly  trans- 
verse. Mandible  usually  small,  with  few  teeth,  ventral  cutting  edge  with  usually  two  or 
three  teeth.  Maxilla  rather  complicated,  of  the  generalized  tipuline  structure.  Antenna 
usually  elongated,  basal  segment  two  to  four  times  as  long  as  it  is  thick,  stouter  in  species 
living  in  decaying  wood.  Mentum  with  seven  to  nine  teeth.  Hypopharynx  a  flattened 
plate,  anterior  margin  usually  with  five  teeth. 

Pupa.- —  Form  generally  stout.  Cephalic  crest  lacking  or  very  small,  with  rudimentary 
setae.  Mouth  parts  as  in  the  subfamily,  sheaths  of  maxillary  palpi  strongly  recurved 
at  tips.  Pronotal  breathing  horns  subequal  in  length,  short,  stout,  usually  straight,  tips 
but  little  expanded.  Mesonotum  transversely  wrinkled,  in  some  wood-inhabiting  species 
(as  T.  trivittata)  with  about  four  conspicuous  tubercles.  Wing  sheaths  and  leg  sheaths 
moderate  in  length.  Abdominal  armature  usually  strong,  each  segment  with  a  posterior 
row  of  four  to  twenty  spines;  in  some  species  a  basal  ventral  row  of  spines  on  posterior  ring 
of  segments.  Cauda  with  dorsal  armature  of  four  powerful  lobes;  eighth  segment  adding, 
as  a  rule,  ten  spines,  of  which  six  are  ventral  and  lateral  in  position,  and  two  or  four  are  dorsal; 
dorso-median  pair  lying  between  anterior  pair  of  lobes  of  cauda,  as  discussed  above,  and 
lacking  or  very  reduced  in  some  wood-inhabiting  species  (T.  trivittata).  Lateral  abdominal 
spiracles  lacking  or  merely  vestigial. 

Tipula  is  the  largest  genus  of  crane-flies,  comprising  a  vast  assemblage 
of  species  (between  six  and  seven  hundred  described  forms)  which  are 
found  on  all  the  continental  areas  of  the  world  but  are  few  in  the  Austral- 
asian region  and  apparently  lacking  on  many  of  the  lesser  oceanic  islands. 
The  genus  is  one  of  extreme  interest,  and  its  study  will  require  many 
years  of  conscientious  application.  Subapterous  species  are  not  rare  in 
this  group,  of  which  many  are  far  northern  forms,  others  are  coastal  spe- 
cies, while  a  few  live  inland  and  under  influences  that  make  it  difficult  to 
explain  their  subapterous  condition. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II 

The  immature  stages  of  the  various  species  are  diverse  in  their  habits, 
ranging  from  spacies  that  are  nearly  if  not  quite  aquatic,  thru  the  majority 
of  the  known  forms  which  live  in  generally  moist  earth  or  mud  along  the 
margins  of  water  bodies,  to  still  others  that  live  in  the  semi-decayed 
wood  of  prostrate  tree  trunks.  Mellor  (1919:64)  has  recorded  Tipula 
larvae  as  breeding  in  manure.  So  far  as  is  known,  the  larvae  are  herbiv- 
orous, tho  they  will  eat  animal  food  under  stress  (as  described  by  Patter- 
son [1908]  for  Tipula  oleracea,  which  feeds  in  considerable  numbers  on 
earthworms) . 

In  Europe,  a  great  number  of  life  histories  in  this  genus  have  been 
worked  out  in  commendable  detail,  mainly  thru  the  efforts  of  Beling, 
who  discusses  no  fewer  than  thirty  species.  His  descriptions  give  a  clear 
idea  of  the  range  in  structure  and  habitat  to  be  expected  in  the  genus. 
The  number  of  lobes  surrounding  the  spiracular  disk  varies  from  four 
(apparently)  in  T.  selene  and  related  forms,  to  as  many  as  eight  in  T. 
subnodicornis.  Practically  all  of  the  known  species  show  the  normal 
tipuline  number  of  lobes,  six. 

A  summary  of  the  larval  habitats  of  the  Palaearctic  species  is  as  follows : 

1.  Species  living  in  saturated  earth  along  watercourses  or  in  debris  at  the  water's  edge, 
or  species  that  are  aquatic  —  Tipula  fulvipennis  de  Geer,  lateralis  Meig.,  lunata  Linn.,  maxima 
Poda,  variicornis  Schum.,  variipennis  Meig.,  vittata  Meig. 

2.  Species  living  in  earth,  usually  in  woods,  underneath  a  mold  of  leaves  or  coniferous 
needles — Tipula  caesia  Schum.,   dilatata  Schum.,  fulvipennis   de  Geer,   hortensis  Meig., 
hortulana  Meig.,  nigra  Linn.,  nubeculosa  Meig.,  ochracea  Meig.,  pabulina  Meig.,  paludosa 
Meig.,  pruinosa  Wied.,  scripta  Meig.,  selene  Meig.,  truncorum  Meig.,  unca  Wied.,  variipennis 
Meig.,  vittata  Meig. 

3.  Species  living  in  earth  in  gardens,  pastures,  or  meadows,  usually  beneath  turf —  Tipula 
irrorata  Macq.,  luteipennis  Meig.,  nigra  Linn.,  ochracea  Meig.,  oleracea  Linn.,  paludosa  Meig., 
pruinosa  Wied.,  subnodicornis  Zett.,  truncorum  Meig.,  vernalis  Meig. 

4.  Species  living  in  or  beneath  cushions  of  moss  or  in  earth  overgrown  with  a  mossy 
covering —  Tipula   dilatata   Schum.,    hortulana   Meig.,   marmorata   Meig.,   pagana   Meig., 
peliostigma  Schum.,  pruinosa  Wied.,  rufina  Meig.,  signata  Staeg.,  truncorum  Meig.,  unca 

5.  Species  living  underneath  moss  on  logs —  Tipula  irrorata  Macq. 

6.  Species  living  in  decaying  wood —  Tipula  flavolineata  Meig.,  irrorata  Macq.,  truncorum 
Meig. 

Bouche*  describes  T.  lunata  and  T.  ochracea  as  living  in  decaying  willow 
wood,  and  Sopotzko  records  T.  flavolineata  as  injuring  clover;  but  these 
records  are  presumably  based  on  mistaken  identifications. 

Comparatively  few  of  the  eastern  American  species  have  been  reared, 
and  it  is  not  considered  advisable  to  attempt  a  key  to  the  larvae  or  the 
pupae  at  this  stage  of  knowledge  of  the  subject.  Such  a  key  would 


1000  CHARLES  PAUL  ALEXANDER 

include  but  a  fraction  of  the  possible  species  and  would  be  of  little  value. 
It  will  require  the  careful  rearing  of  species  for  many  years  before  a 
workable  key  to  the  immature  stages  of  the  eastern  species  of  the  genus 
can  be  produced.  The  characters  that  will  prove  of  greatest  value  in 
the  separation  of  the  larvae  and  the  pupae  of  the  species  of  Tipula  are  as 
follows: 

Larvae 

1.  Anal  gills.     (These  are  rarely  lacking,  and  the  number  and  arrangement  of  the  branches, 
their  form,  and  their  function,  are  of  primary  importance.) 

2.  Spiracular  disk.     Number  of  lobes  surrounding  disk  and  whether  they  are  simple  or 
branched;  character  and  nature  of  fringe  of  hairs  around  disk,  if  such  is  present;  size,  shape, 
and  distance  apart  of  spiracles;  markings  on  inner  face  of  disk  and  lobes. 

3.  Chaetotaxy.     Arrangement,  length,  and  number  of  setae  on  segments. 

4.  Body  form.    Terete,  subdepressed,  or  flattened  ventrally  only;  clothing  of  pubescence, 
and  pattern  formed  on  dorsum. 

5.  Head  capsule.     (The  head  is  remarkably  uniform  thruout  the  group,  a  condition  to 
be  expected  in  a  group  so  compact  as  Tipula.)     Shape  of  mentum  and  hypopharynx,  and 
number,  size,  and  shape  of  teeth  along  their  anterior  margins;  shape  of  antenna,  and  other 
details  of  head. 

Pupae 

1.  General  form,  whether  terete  or  depressed. 

2.  Mouth  parts. 

3.  Pronotal  breathing  horns,  their  relative  length,  size,  and  form. 

4.  Armature  of  mesonotum. 

5.  Wing  sheaths  and  leg  sheaths. 

6.  Spines  on  abdominal  segments,  their  size  and  number;  whether  lacking  or  present  at 
base  of  posterior  ring  of  sternites;  arrangement  and  number  of  pleural  spines. 

7.  Cauda,  shape  of  genital  sheaths,  armature  of  dorsum,  and  ventral  margin  of  eighth 
eternite. 

Descriptions  are  given  in  the  following  pages  of  about  ten  life  histories 
which  are  entirely  new  or  have  been  insufficiently  considered  elsewhere. 
A  few  notes  on  certain  other  species  that  have  been  observed  in  the  past 
few  years  may  be  added  here: 

Tipula  cayuga  Alex.  A  conspicuous  yellow  larva,  living  in  organic  earth  beneath  leaves, 
in  association  with  Bittacomorphella  jonesi  and  other  forms  which  are  discussed  elsewhere 
(page  781).  The  pupal  duration  is  slightly  over  seven  days. 

T.  angustipennis  Loew.  Found  living  in  rather  dry  earth  beneath  leaves  in  shaded  woods 
(Lawrence,  Kansas,  Mrs.  C.  P.  Alexander). 

T.  umbrosa  Loew.  Occurs  in  garden  soil  in  company  with  the  larvae  of  Tipula  bicornis 
Forbes. 

T.  fuliginosa  Say.  Reared  from  larvae  living  in  debris  under  the  nest  of  a  turkey  vulture 
(Jackson  Island,  Maryland,  May  23,  1913,  R.  C.  Shannon). 

T.  sayi  Alex,  and  T.  tricolor  Fabr.     In  saturated  mud  in  marshy  or  swampy  situations. 

2T.  tephrocephala  Loew.     A  large  larva,  nearly  if  not  quite  aquatic  in  its  habits. 


THE  CRANE-FLIES  OF  NEW  YORK  — PART  II  1001 

The  life  histories  of  other  Nearctic  species  are  recorded  in  the  sum- 
mary of  literature  on  page  980.  These  are  as  follows:  Tipula  arctica 
Curt.  (Nielsen,  Alexander),  T.  eluta  Loew  (Hart,  Malloch),  T.  cunctans 
Say  (Hyslop,  Malloch),  T.  bicornis  Forbes  (Forbes),  and  T.  ultima  Alex. 
(Needham,  Caudell).  T.  arctica,  according  to  Nielsen  (1910:57-59),  was 
found  commonly  in  eastern  Greenland.  The  immature  stages  were  dis- 
covered in  circular  holes  from  two  to  three  centimeters  deep  in  the  ground, 
especially  beneath  tufts  of  Cassiope  tetragona  (L.)  D.  Don.  The  pupae 
were  found  at  the  end  of  June,  and  empty  pupa  cases  were  found  as  early 
as  the  25th  of  the  same  month.  According  to  Nielsen,  the  larvae  require 
two  years  to  attain  their  growth. 

Tipula  (Trichotipula)  oropezoides  Johns. 

1909     Tipula  oropezoides  Johns.    Proc.  Boston  Soc.  Nat.  Hist.,  vol.  34,  p.  131-132. 

Larvae  of  Tipula  oropezoides  were  first  found  on  March  30,  1917,  living 
beneath  saturated  moss  in  Needham Js  Glen,  Ithaca,  New  York,  where 
they  were  associated  with  larvae  of  Dicranomyia  badia,  Penthoptera 
albitarsis,  Tipula  collaris,  and  other  species.  Numerous  additional  larvae 
were  found  in  the  same  locality  on  April  18,  1917.  Some  of  these  were 
placed  in  rearing  and  emerged  on  May  6.  The  larvae  are  nocturnal  in 
their  habits,  being  very  sluggish  and  retiring  during  the  day  but  becoming 
active  after  sunset. 

The  adult  flies  bear  a  strong  resemblance  to  species  of  the  genus  Oropeza, 
with  which  they  are  sometimes  found  associated.  They  may  often  be 
swept  from  rank  herbage  in  cool  Canadian  woods. 

Larva. —  Length,  16.5-17  mm. 
Diameter,  1.8-2  mm. 

Coloration  above,  a  deep  velvety  brown  with  mottlings  of  paler;  on  basal  ring  of  tergites 
six  median  transverse  pale  spots,  posterior  ring  less  regularly  marked;  pleura  and  venter 
pale.  (The  dark  markings  on  the  dorsum  are  produced  by  patches  of  dark-colored  hairs, 
which  cover  the  body  densely  in  places.) 

Form  subterete.  Integument  with  an  abundant  pubescence,  longest  on  dorsum.  Chaeto- 
taxy  as  follows:  dorsum  (Plate  XCIII,  516)  on  posterior  ring  with  six  stout  setae,  three  on 
either  side,  the  middle  seta  a  little  closer  to  the  inner  seta;  ventral  segments  (Plate  XCIII, 
517)  with  four  setae,  two  anterior  and  two  posterior,  the  latter  a  little  more  separated. 
Spiracular  disk  (Plate  XCIII,  518)  surrounded  by  six  approximately  subequal  lobes,  their 
inner  faces  heavily  lined  with  dark  brown;  at  tip  of  each  lobe  a  pale  rounded  spot,  largest  on 
ventral  lobes  and  here  with  a  sensory  bristle;  lateral  mark  not  reaching  spiracles;  lateral 
and  dorsal  lobes  slightly  paler  medially;  above  and  below  each  spiracle  a  transverse  brown 


1002  CHARLES  PAUL  ALEXANDER 

line;  on  disk,  between  spiracles,  two  indistinct  dusky  spots;  lobes  fringed  with  rather  short, 
pale  hairs  which  are  narrowly  interrupted  between  lobes.  Spiracles  irregular,  roughly 
triangular.  Anal  gills  four,  slender,  posterior  pair  the  larger  (Plate  XCII,  506). 

Head  capsule  as  in  genus.  Labrum  and  maxilla-very  densely  fringed  with  long  golden- 
yellow  hairs.  Mentum  (Plate  XCIII,  513)  with  two  flattened  lateral  teeth,  the  median 
point  elongated;  mentum  very  deeply  split  behind.  Hypopharynx  (Plate  XCIII,  514)  with 
but  three  evident  teeth,  the  lateral  teeth  very  broad,  flattened.  (In  some  specimens  these 
teeth  are  all  very  blunt,  so  that  the  anterior  margin  of  the  hypopharynx  appears  merely 
crenulate.)  Antenna  with  apical  disk  very  flattened.  Mandible  (Plate  XCIII,  515)  with 
a  dorsal  tooth  and  a  powerful  ventral  tooth. 

Pupa. —  Length:  male,  12  mm.;  female,  12.5-13  mm. 

Width,  d.-s.:  male,  1.6-1.7  mm.;  female,  1.8-1.9  mm. 
Depth,  d.-v.:  male,  1.6  mm.;  female,  1.7-1.8  mm. 

Coloration  dark  brown;  dorsum  of  thorax  and  abdomen,  and  face,  more  reddish  brown. 

General  features  as  in  Tipula  collaris.  Form  slender.  Pronotal  breathing  horns  long 
and  slender,  dark-colored,  divergent  at  tips.  Antenna  elongate.  Wing  sheaths  ending 
opposite  apex  of  second  abdominal  segment.  Leg  sheaths  long,  extending  to  beyond  mid- 
length  of  fourth  abdominal  segment.  Male  cauda  with  dorsal  lobes  of  genitalia  short,  blunt; 
ventral  lobes  produced  caudad  into  slender,  blunt  lobes  which  are  transversely  wrinkled, 
separated  by  a  U-shaped  notch,  at  base  on  outside  with. a  prominent  spine.  Female  ovi- 
positor elongate;  dorsal  valves  narrowed  to  the  blunt  tip;  ventral  lobes  stout,  a  little  shorter 
than  dorsal  valves,  tips  strongly  divergent;  the  six  dorsal  lobes  of  cauda  spinous-tipped, 
sharply  pointed. 

Nepionotype  —  Ithaca,  New  York,  April  18,  1917.     No.  6-1917. 
Neanotype. —  With  type.     No.  7  1917. 
Paratypes. —  Numerous  larvae  and  pupae. 

Tipula  collaris  Say 

1823     Tip-da  collaris  Say.    Journ.  Acad.  Nat.  Sci.  Phila.,  vol.  3,  p.  23. 

Larvae  and  pupae  of  Tipula  collaris,  a  common  vernal  crane-fly, 
occurred  frequently  beneath  saturated  moss  (Amblystegium  irriguum 
[Wils.]  B.  &  S.)  in  Needham's  Glen,  Ithaca,  New  York,  on  April  17, 
1917.  Their  associates  are  noted  under  the  •  account  of  T.  oropezoides 
(page  1001).  In  the  same  moss  areas  occurred  numerous  small  red-backed 
salamanders  (Plethedon  cinereus),  which  probably  fed  on  the  insect  deni- 
zens of  the  place.  Specimens  emerged  in  the  writer's  breeding  jars  as  late 
as  May  10. 

The  adults  are  on  the  wing  during  April  and  May,  some  persisting  into 
early  June  in  cool  northern  woods.  The  life  history  undoubtedly  requires 
a  year  for  its  completion. 

Larva. —  Length,  21.5-25  mm. 
Diameter,  2.5-3.5  mm. 


THE  CRANE-FLIES  OF  NEW  YORK — PART  II  1003 

Coloration  pale  dusky,  darker  above;  dorsum  behind  with  two  broken  lines  which  are 
divergent  on  each  annulus,  those  of  anterior  annulus  made  up  of  three  circular  spots;  pleura 
with  a  conspicuous  dark  brown  stripe;  venter  almost  uniformly  pale,  with  indistinct  lines; 
anterior  part  of  thoracic  segments  darker.  (In  life  the  color  is  rich  reddish  brown,  and  the 
pleural  stripe  is  not  evident.) 

Form  terete.  Body  covered  with  a  delicate,  pale  pubescence,  in  addition  to  the  usual 
setae.  Chaetotaxy  as  follows:  tergites  (Plate  XCIV,  521)  with  a  transverse  row  of  six  setae 
on  posterior  ring  before  margin,  the  two  innermost  solitary,  each  lateral  pair  closely 
approximated;  a  solitary  seta  on  extreme  lateral  margins  of  tergite,  at  margin  of  dark  pleural 
stripe,  and  at  about  midlength  of  posterior  ring;  pleura  on  basal  ring  with  a  single  seta,  pos- 
terior ring  with  a  group  of  about  three  or  four  setae,  one  larger  than  the  others;  sternites 
with  four  widely  separated  setae  on  posterior  ring,  the  lateral  pair  a  little  nearer  posterior 
margin  than  the  median  pair.  Spiracular  disk  (Plate  XCIV,  522)  surrounded  by  six  lobes 
which  are  moderately  slender;  ventral  pair  a  little  longer,  dorsal  pair  a  little  shorter;  all  the 
lobes  broadly  margined  with  pale  brown;  ventral  lobes  having  in  addition  a  black  capillary 
line  extending  from  tips  backward  to  beyond  midlength  of  lobes,  this  line  broadest  at  tip, 
gradually  narrowing,  and  becoming  paler  toward  base  of  lobes;  below  each  spiracle,  two 
conspicuous  black  dots;  lobes  fringed  with  numerous  rather  long  hairs.  Spiracles  large, 
separated  by  a  distance  a  little  greater  than  diameter  of  one.  Anal  gills  six,  one  pair  much 
shorter  than  the  others,  consisting  of  a  basal  branch  of  the  anterior  gill. 

Head  capsule  rather  small,  of  the  usual  tipuline  type.  Mentum  (Plate  XCIV,  519)  very 
broad,  anterior  margin  almost  transverse,  seven-toothed,  median  point  the  longest.  Hypo- 
pharynx  (Plate  XCIV,  520)  with  five  teeth,  which  are  very  short  and  blunt  giving  anterior 
margin  a  deeply  crenulated  appearance;  before  hypopharynx  a  rounded  lobe  which  is  densely 
covered  with  six  short,  blunt,  chitinized  points,  this  being  probably  the  prementum.  Antenna 
elongate-cylindrical;  apex  with  apical  disk  very  small,  button-like.  Mandible  small,  with 
about  one  dorsal  and  two  ventral  teeth.  Maxilla  slender,  densely  hairy;  palpus  subglobular, 
with  several  small,  hyaline  papillae. 

Pupa. —  Length,  17-19  mm. 

Width,  d.-s.,  2.8-3.2  mm. 
Depth,  d.-v.,  3-3.3  mm. 

Coloration  brown;  wing  sheaths,  except  in  older  individuals,  pale;  pleural  region  of  abdomen 
light  yellow;  abdominal  incisures  often  pale. 

Head  rather  small.  Cephalic  crest  low  and  indistinct,  with  tiny  setae.  Labrum  broad, 
apex  pointed.  Labial  lobes  oval,  contiguous  at  inner  end.  Maxillary  palpi  strongly  recurved 
at  tips.  Antenna  slender,  moderately  elongated,  extending  some  distance  beyond  wing 
root.  Pronotal  breathing  horns  equal  in  length,  rather  short,  the  moderately  long  tips  flat- 
tened, smooth;  two  small,  approximated  setae  on  either  side  of  median  line.  Wing  sheath 
(Plate  XCIV,  523)  extending  just  beyond  end  of  second  abdominal  segment;  venation  dis- 
tinct. Leg  sheaths  extending  beyond  base  of  fourth  abdominal  segment;  fore  tarsi  con- 
siderably shorter  than  the  others. 

Abdominal  segments  with  armature  of  posterior  ring  weak,  the  spines  very  short  and  stout, 
with  a  few  setae;  maximum  number  of  spines  on  the  tergites  about  twenty;  lateral  anterior 
angle  of  posterior  ring  of  tergites  with  two  small  spines:  pleurites  with  one  basal  spine,  and 


1004  CHARLES  PAUL  ALEXANDER 

three  setiferous  spines  on  posterior  ring,  arranged  transversely;  spiracles  very  rudimentary, 
opposite  base  of  posterior  ring;  sternites  with  spines  slightly  more  numerous  and  stouter; 
near  base  of  posterior  ring  two  transverse  spines  on  either  side  median  line,  the  outermost 
setiferous.  Male  cauda  on  dorsum  with  six  lobes;  the  four  posterior  ones  stout,  with  tips 
spinous  or  those  of  ventral  pair  slightly  bifid;  the  two  anterior  median  lobes  shorter  and 
more  slender;  eighth  sternite  with  four  large  spines  about  equally  spaced;  eighth  pleurite 
with  a  large,  powerful,  acutely  tipped  lobe,  and  two  or  three  smaller  dorso-lateral  spines 
above  base  of  middle  pair  of  dorsal  lobes.  Female  cauda  almost  the  same  as  male  cauda, 
due  to  the  blunt  terebral  sheaths  of  this  species. 

Nepionotype.—  Ithaca,  New  York,  March  30,  1917. 

Neanotype. —  With  type  larva. 

Paratypes. —  Numerous  larvae  and  pupae,  March  30  to  April  18,  1917. 

Tipula  nobilis  (Loew) 

1864     Pachyrrhina  nobilis  Loew.    Berl.  Ent.  Ztschr.,  vol.  8,  p.  62. 

Larvae  of  Tipiula  nobilis  were  found  in  wet  moss  and  beneath  decaying 
witch-hazel  leaves  at  Orono,  Maine,  on  June  17,  1913.  An  adult  emerged 
on  July  1.  A  fully  grown  pupa  found  on  July  11  attempted  to  transform, 
but  died  after  two  hours  without  being  able  to  extricate  itself  from  the 
pupal  case. 

The  adults,  which  strikingly  resemble  some  species  of  Nephrotoma,  fly 
somewhat  later  than  does  T.  collaris,  but  both  species  may  be  taken 
together  in  early  June. 

T.  nobilis  is  very  similar  in  all  respects  to  T.  collaris. 

Larva. —  Length,  20  mm. 

Diameter,  2.9-3  mm. 

Coloration,  reddish  brown. 

Spiracular  disk  as  in  T.  collaris,  the  brown  lateral  margin  to  the  lobes  a  little  paler.  Anal 
gills  as  shown  in  Plate  XCII,  507. 

Head  capsule  almost  as  in  T.  collaris.  Mentum  with  apical  point  elongate,  with  three 
blunt  teeth  on  either  side.  Hypopharynx  with  five  moderately  acute  teeth.  Antenna  with 
a  blunt  conical  papilla,  larger  and  more  conspicuous  than  in  T  coUaris.  Mandible  with  teeth 
very  blunt. 

Pupa. —  Length  of  cast  pupal  skin,  about  18.5  mm. 

Pupa  very  similar  to  that  of  T.  collaris. 

Nepionotype.—  Orono,  Maine,  June  19,  1913.    No.  40-1913. 

Neanotype. —  With  type. 

Tipula  bella  Loew 

1863    Tipula  betta  Loew.    Berl.  Ent.  Ztechr.,  vol.  7,  p.  291-292. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  1005 

Tipula  bella  is  a  common  species,  flying  thruout  the  summer.  Larvae 
and  pupae  are  not  rare  in  sandy  or  loamy  soil  along  streams.  A  larva 
taken  on  April  28,  1917,  in  gravel  at  Ithaca,  New  York,  where  it  was 
associated  with  larvae  of  Hexatoma,  was  placed  in  rearing.  It  emerged 
as  an  adult  male  on  May  15.  On  May  27,  1913,  three  fully  colored  pupae 
were  found  along  the  sandy  banks  of  Fall  Creek,  Ithaca,  in  association 
with  Eriocera  spinosa.  They  emerged  as  females  on  May  29  and  30. 

Larva. —  (The  description  is  from  field  notes  on  the  larva  mentioned  above.) 
Length,  20-25  mm. 

Coloration  light  grayish  brown,  with  a  slight  reddish  cast  most  noticeable  on  venter; 
dorsum  with  two  narrow,  almost  continuous,  dark  brown  lines,  these  lines  subparallel  at 
anterior  part  of  each  segment,  then  strongly  bellied  out,  and  then  parallel  but  finally 
divergent. 

Spiracular  disk  surrounded  by  six  rather  short  lobes;  ventral  lobes  with  a  linear,  rather 
pale,  brown  mark,  and  a  few  sensory  bristles  at  tips;  lateral  lobes  with  ventral  margin  lined 
with  brown;  dorsal  lobes  with  both  margins  feebly  bordered  with  brown;  two  small  brown 
dots  below  each  spiracle.  Anal  gills  six,  very  long  and  slender. 

Pupa.  —  Length  of  cast  pupal  skin,  about  24  mm. 

Pupa  similar  to  pupae  of  other  species  of  genus.  Pronotal  breathing  horns  short, 
cylindrical,  narrowed  to  tips.  Spines  on  abdominal  segments  rather  large,  especially  on 
sternites;  spines  on  base  of  posterior  ring  of  sternites  small  but  evident.  Female  cauda 
with  dorsal  valves  long,  pointed;  sternal  valves  shorter.  Cauda  with  the  usual  six  lobes 
on  dorsum,  the  four  posterior  stout,  divergent,  spinous-tipped,  the  anterior  median  pair 
much  smaller;  venter  of  segment  8  with  three  strong  spines  on  either  side,  gradually  smaller 
from  lateral  spine  toward  innermost  spine. 

Neanotype. —  Ithaca,  New  York,  reared  May  25,  1917. 

Paratypes  —  Pupal  skins,  type  locality,  May  5,  1914  (No.  42-1914);  August,  1911;  May 
25,  1917;  etc. 

Tipula  caloptera  Loew 

1863     Tipula  caloptera  Loew.     Berl.  Ent.  Ztschr.,  vol.  7,  p.  292. 

The  vigorous  larva  of  Tipula  caloptera  is  one  of  the  largest  and  most 
striking  in  the  family.  The  larvae  live  in  rapid-  or  slow-flowing  streams 
either  in  the  water  among  debris  and  under  stones,  or  in  the  sand,  gravel, 
or  mud  in  very  close  proximity  to  the  water.  Here  they  are  associated 
with  the  larvae  of  various  species  of  Eriocera,  Erioptera  armata,  Tabanus, 
Atherix,  and  other  forms.  Oftentimes  they  are  found  in  deep  water  in 
exceedingly  lotic  situations.  A  larva  placed  in  rearing  on  April  19,  1917, 
emerged  as  an  adult  female  on  May  13. § 


1006  CHARLES  PAUL  ALEXANDER 

This  is  evidently  the  larva  taken  by  Dr.  Needham  in  the  Adirondacks 
and  referred  by  him  with  some  doubt  (Needham  and  Betten,  1901:575- 
576)  to  T.  abdominalis.  This  dubious  reference  has  created  considerable 
confusion  ever  since  the  species  was  figured  on  the  cover  of  Entomological 
News  under  the  facetious  name  "  Quisnam  sexcaudatusf  "  Malloch  (1915- 
17  b:  200-201)  mentions  the  same  larva  under  the  name  T.  abdominalis. 
As  stated  elsewhere,  the  larva  of  abdominalis  is  very  different. 

Larva. —  Length,  45-55  mm. 

Diameter,  4.8-6.8  mm. 

Coloration  above,  dark  brown  or  brownish  green;  segments  beautifully  marked  with  small 
white  spots,  especially  anterior  segments;  a  broad,  dark  brown,  median  stripe,  and  a  more 
or  less  distinct  pale  lateral  stripe  (in  preserved  specimens  the  pleura  is  usually  dark  brown, 
more  distinct  behind,  with  numerous  pale  white  dots);  sternum  dark  greenish.  (In  older 
specimens  the  color  is  very  dark  and  the  pattern  is  more  or  less  obliterated.) 

Form  stout,  terete.  Body  smooth,  segments  with  indistinct  posterior  tubercles.  Chaeto- 
taxy  very  weak,  a  few  weak  pleural  setae  on  posterior  ring,  sternal  and  tergal  setae  minute. 
Spiracular  disk  rather  small,  surrounded  by  six  subequal,  moderately  narrow,  lobes  which 
are  fringed  with  short  hairs;  margins  of  disk  and  lobes  somewhat  as  in  T.  bella,  each  lobe  with 
a  delicate  capillary  brown  line;  two  brown  spots  beneath  each  spiracle.  Spiracles  small, 
separated  by  a  distance  about  equal  to  twice  diameter  of  one.  Anal  gills  six,  very  long  and 
slender  (Plate  XCII,  508). 

Head  capsule  as  in  genus.  Men  turn  broad;  anterior  margin  nearly  transverse,  with  three 
subacute  teeth  on  either  side,  median  point  not  conspicuously  elongated. 

Pupa.- —  Length  of  cast  skin,  about  32-35  mm. 

Characters  almost  as  in  T.  bella.  Pronotal  breathing  horns  short,  cylindrical,  tips  not 
expanded.  Abdominal  spines  prominent,  projecting,  few  in  number,  on  intermediate  ter- 
gites  9  or  10;  pleurites  with  a  single  strong  spine  on  each  ring;  sternites  with  a  posterior  row 
of  seven  or  eight  strong  spines;  on  segments  5  to  7  two  strong  spines  at  base  of  posterior 
ring,  those  of  seventh  segment  the  largest.  Cauda  almost  as  in  T.  bella. 

Nepionotype  —  Ithaca,  New  York,  April  26,  1917.     No.  16-1917. 
Neanotype. —  Cast  pupal  skin,  reared  May  13,  1917. 
Paratypes. —  Larvae  and  cast  pupal  skins  from  type  locality. 

Tipula  dejecta  Walk. 

1856     Tipula  dejecta  Walk.     Ins.  Saunders,  vol.  1,  Dipt.,  p.  442. 
1901     Tipula  fumosa  Doane.     Journ.  N.  Y.  Ent.  Soc.,  vol.  9,  p.  99. 

Tipula  dejecta  is  a  characteristic  vernal  species  flying  in  April  and  May. 
The  flies  are  notable  inhabitants  of  swamps,  especially  alder  swamps. 

On  April  20,  1917,  the  writer  found  larvae  of  this  fly  in  Larch  Meadows, 
near  Ithaca,  New  York,  in  association  with  larvae  of  Rhamphidia  main- 
ensis,  Pseudolimnophila  luteipennis,  and  other  swamp  inhabitants.  The 


THE  CRANE-FLIES  OF  NEW  YORK  — PART  II  1007 

conditions  are  discussed  more  fully  under  the  account  of  Rhamphidia 
(page  831).  The  larvae  are  reddish  brown  in  color,  and  rather  sluggish. 
One  of  the  larvae  found  on  April  20  pupated  on  the  22d  and  emerged  as 
an  adult  female  on  the  30th,  a  pupal  duration  of  eight  days.  An  additional 
pupa  was  taken,  associated  with  larva  of  Pseudolimnophila  luteipennis, 
P.  inornata,  Tricyphona  inconstans,  and  other  species. 

Larva. —  Length,  20  mm. 

Diameter,  1.8  mm. 

Coloration  brown;  dorsum  marked  with  light  and  dark  brown;  a  narrow,  indistinct,  dark 
brown,  median  line,  with  a  broader  zigzag  brown  line  on  either  side;  ventral  surface  a  little 
paler. 

Body  covered  with  a  short,  dark  pubescence  at  sides  of  segments,  at  margins  longer  and 
more  conspicuous.  Chaetotaxy  as  follows:  tergites  with  six  strong  setae  in  transverse 
alinement,  the  outermost  in  pairs;  two  strong  setae  on  each  pleural  annulus;  posterior  ring 
of  sternites  with  eight  strong  setae,  arranged  in  four  pairs.  Spiracular  disk  (Plate  XCV, 
526)  pale,  surrounded  by  six  approximately  equal  lobes  which  are  heavily  marked  with 
brown;  dorsal  and  lateral  pairs  pointed,  ventral  pair  blunt;  ventral  lobes  with  apical  half 
shiny  black,  on  ventral  inner  margin  continued  dorsad,  almost  contiguous  on  midline;  inner 
face  of  dorsal  and  lateral  lobes  suffused  with  dark  brown,  proximal  margin  of  dorsal  lobe 
produced  inward  so  that  the  marks  are  almost  contiguous  on  median  line;  beneath  each 
spiracle  a  transversely  rectangular,  dark  brown  mark.  Anal  gills  with  four  anterior  lobes 
which  are  long  and  slender,  and  a  pair  of  rudimentary  blunt  posterior  gills  (Plate  XCII,  509). 

Head  capsule  and  mouth  parts  as  in  genus.  Men  turn  (Plate  XCV,  524)  seven-toothed, 
apical  point  the  longest.  Hypopharynx  (Plate  XCV,  525)  bluntly  five-toothed. 

Pupa. —  Length,  15.3  mm. 

Width,  d.-s.,  2.3-2.4  mm. 
Depth,  d.-v.,  2.1-2.2  mm. 

Coloration  dark  brown;  abdominal  incisures  paler. 

Form  relatively  stout.  General  features  as  in  genus.  Cephalic  crest  consisting  of  two 
blunt  lobes  with  microscopic  setae.  Maxillary  palpi  strongly  curved  at  tip,  but  not  entirely 
recurved.  Pronotal  breathing  horns  short,  tips  a  little  enlarged. 

Abdominal  tergites  with  spines  weak,  on  median  area  of  each  row  weak  or  lacking;  on 
intermediate  segments  about  fifteen  spines;  pleurites  with  only  a  single  weak  spine  on  basal 
ring;  on  posterior  ring  a  rudimentary  anterior  spine  and  a  somewhat  larger  posterior  spine; 
sternites  similarly  armed  to  tergites,  but  spines  fewer  in  number  and  larger,  on  segment 
5  about  twelve  in  number;  on  base  of  posterior  ring  a  large  spine  on  either  side  median  line 
and  a  small  setiferous  tubercle  laterad  of  each.  Female  cauda  with  tergal  valves  of  ovi- 
positor long  and  straight,  sternal  valves  a  little  shorter;  cauda  with  the  usual  six  dorsal  lobes, 
these  terminating  in  slender  spines;  at  end  of  eighth  sternite  six  large  spines;  dorsal  spines 
reduced  to  a  single  small  pair,  one  near  each  lateral  margin. 

Nepionotype  —  Larch  Meadows,  Ithaca,  New  York,  April  20,  1917. 

Neanotype. —  With  type. 

Paratypes. —  Two  pupae  with  type  pupa. 


1008  CHARLES  PAUL  ALEXANDER 

Tipula  usitata  Doane 

1901     Tipula  usitata  Doane.    Journ.  N.  Y.  Ent.  Soc.,  vol.  9,  p.  124. 

A  large  number  of  fully  grown  larvae  of  an  unknown  species  of  Tipula 
were  found  beneath  the  bark  of  a  fallen  tree  at  Stanford  University, 
California,  on  March  22,  1915,  by  Harold  Morrison.  They  were  sent 
to  the  writer  at  Ithaca,  New  York,  and  emerged  as  adults  on  April  15. 
The  immature  stages  are  very  distinct,  closest  perhaps  to  T.  trivittata, 
which  also  lives  beneath  the  bark  of  decaying  trees.  Nothing  is  known 
of  the  habits  of  the  adult  flies. 

Larva. —  Length,  25-27  mm. 
Diameter,  3-3.2  mm. 

Coloration  pale  greenish  yellow,  darker  above. 

Form  terete.  Body  with  a  very  sparse  pubescence.  Chaetotaxy  as  follows:  tergites 
(Plate  XCV,  527)  with  a  posterior  row  of  eight  setae,  the  middle  pair  of  each  side  very  closely 
approximated;  a  seta  near  lateral  margin  at  base  of  posterior  ring,  on  a  level  with  pleural 
seta;  pleurites,  one  seta  on  each  ring;  sternites  with  eight  setae  in  closely  approximated  pairs 
on  posterior  ring.  Spiracular  disk  (Plate  XCV,  528)  surrounded  by  six  lobes;  dorsal  and 
lateral  pairs  slender,  tips  of  former  acute;  ventral  lobes  blunt;  ventral  lobes  with  tips  black- 
ened, continued  down  proximal  margin  of  lobes  as  a  paler  brown  line;  lateral  lobes  with  inner 
face  narrowly  blackened,  this  mark  not  reaching  spiracles;  dorsal  lobes  with  entire  inner  face 
bulging,  intensely  black,  the  marks  contiguous  at  their  basal  inner  angle;  an  indistinct  brown 
spot  underneath  each  spiracle,  in  some  specimens  this  mark  continuous  with  that  of  ventral 
lobes;  lobes  not  fringed  with  hairs.  Spiracles  large,  separated  by  a  distance  about  equal  to, 
or  a  little  greater  than,  diameter  of  one.  Anal  gills  short  and  blunt,  strongly  protuberant, 
surrounding  anus  as  four  fleshy  lobes  (Plate  XCII,  510). 

Head  capsule  as  in  genus.  Mentum  broad,  with  seven  to  nine  teeth,  in  the  latter  case 
the  outermost  pair  very  small.  Hypopharynx  with  three  or  five  very  blunt  teeth.  Antenna 
much  shorter  and  stouter  than  in  most  species  of  Tipula,  the  length  only  a  little  greater  than 
twice  the  diameter,  at  apex  with  a  blunt  conical  papilla  and  a  few  small,  cylindrical  sense 
pegs.  Mandible  powerful,  with  two  or  three  flattened  teeth  on  ventral  cutting  edge. 

Pupa. —  Length,  15-16.8  mm. 
Width,  d.-s.,  2  mm. 
Depth,  d.-v.,  2.3-2.4  mm. 

Coloration  pale  brown;  posterior  margin  of  abdominal  rings  pale;  lateral  margin  of 
abdomen  conspicuously  pale  yellowish  white. 

Form  slender.  Pronotal  breathing  horns  narrow,  a  little  expanded  at  tips.  Leg  sheaths 
ending  on  a  level. 

Abdominal  spines  very  strong,  but  few  in  number;  tergites  with  four  to  six  spines;  pleurites 
with  a  single  weak  spine  on  each  ring;  sternal  spines  very  strong,  five  or  six  in  number,  those 
on  segment  7  subequal  in  size  to  those  on  segment  8;  no  sternal  spines  on  base  of  posterior 
ring.  Male  cauda  with  posterior  dorsal  lobe  very  strong,  pale,  tips  acute;  lateral  lobes  greatly 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  1009 

reduced  in  size,  the  median  pair  represented  only  by  two  blunt  brown  tubercles;  eighth 
segment  with  the  usual  six  strong  ventral  and  lateral  lobes.  Female  ovipositor  with  valves 
small;  sternal  valves  short  and  broad,  much  shorter  than  tergal  valves. 

Nepionotype. —  Stanford  University,  California,  April  15,  1915. 
Neanotype. —  With  type,  bred  April  15,  1915. 
Paratypes. —  Numerous  larvae  and  pupae  with  type. 

Tipula  trivittata  Say 

1823     Tipula  trivittata  Say.     Journ.  Acad.  Nat.  Sci.  Phila.,  vol.  3,  p.  26. 

The  larvae  and  the  pupae  of  Tipula  trivittata  live  beneath  the  bark  of 
much-decayed  prostrate  trunks  or  under  the  layers  of  moss  that  often 
cover  fallen  trees.  Abundant  larvae  were  found  on  March  22,  1913,  and 
were  placed  in  rearing,  adults  emerging  on  April  26.  On  April  16,  1914, 
larvae  of  two  distinct  sizes  — some  very  small  and  some  nearly  fully  grown 
—  occurred  in  abundance  beneath  moss  (Entodon  seductrix  [Hedw.]  C. 
MuelL,  Brachythecium  acuminatum  [Hedw.]  Kindb,,  Hypnum  Haldan- 
ianum  Grev.,  and  Mnium  sylvaticum  Lindb.)  on  prostrate  decaying  elms, 
sycamores,  and  other  trees,  at  Renwick  Park,  Ithaca,  New  York. 

The  adult  flies  are  among  the  commonest  of  the  eastern  specjes  of 
Tipula,  and  fly  during  a  large  part  of  the  season. 

Larva. —  Length,  24-25  mm. 
Diameter,  2.7-3  mm. 

Coloration  pale  brownish  yellow,  a  little  paler  beneath. 

Form  moderately  elongated,  terete.  Pubescence  very  short  or  practically  lacking.  Setae 
on  anterior  segments  strong,  on  posterior  segments  shorter.  Chaetotaxy  as  follows:  ter- 
gites  with  a  posterior  row  of  six  setae,  the  two  middle  punctures  each  with  a  single  seta, 
the  two  lateral  punctures  each  with  two  setae;  pleurites  with  a  single  seta  on  each  ring; 
sternites  with  two  rows  of  setae,  the  anterior  row  consisting  of  two  closely  approximated 
groups  of  two  setae  each,  the  posterior  row  consisting  of  a  single  large  seta,  laterad  of  which 
is  a  minute  bristle.  Spiracular  disk  (Plate  XCV,  529)  surrounded  by  six  lobes;  lateral  pair 
long  and  slender;  dorsal  pair  a  little  shorter,  slender;  ventral  lobes  blunt;  ventral  lobes  with 
a  jet-black  mark  on  inner  face;  lateral  lobes  with  dark  markings  represented  only  by  a  very 
small  linear  dash;  dorsal  lobes  with  a  small  black  area.  Spiracles  large,  separated  by  a  dis- 
tance a  little  less  than  diameter  of  one.  Anal  gills  indistinctly  lobed,  four  in  number,  two 
on  either  side,  very  blunt  and  protuberant. 

Head  capsule  as  in  genus,  the  mouth  parts  almost  as  in  T.  usitata.  Mentum  broad,  with 
seven  teeth.  Hypopharynx  with  five  teeth,  the  three  middle  ones  the  longest,  subequal 
in  size.  Antenna  shorter  and  stouter  than  is  usual  in  the  genus. 

Pupa. —  Length,  19-20  mm. 

Width,  d.-s.,  2.5-2.6  mm. 
Depth,  d.-v.,  2.8-3  mm. 

11 


1010  CHARLES  PAUL  ALEXANDER 

Coloration  dark  brown;  abdomen  yellow,  with  a  broad  sublateral  brown  stripe  on  both 
sternites  and  tergites;  abdominal  segments  beyond  posterior  row  of  spines  brighter,  more 
yellowish. 

Characters  of  head  as  in  genus.  Cephalic  crest  very  small.  Labrum  broad.  Pronotal 
breathing  horns  short,  slightly  curved.  Mesonotal  prescutum  with  fine  transverse  wrinkles; 
two  blunt  lateral  tubercles,  and  behind  these,  on  either  side  of  median  line,  two  smaller 
flattened  ledges  which  are  often  bifid  at  their  tips.  (Similar  ledges,  but  much  less  prominent, 
occur  in  T.  usitata.} 

Abdominal  tergites  with  subapical  armature  weak,  spines  varying  in  number  from  six  to 
eight,  those  of  posterior  segments  larger;  pleural  spines  long  and  slender,  one  on  each  ring; 
sternal  spines  powerful,  four  to  six  in  number,  no  spines  on  base  of  posterior  ring.  Female 
cauda  with  sternal  valves  long  and  slender,  but  little  shorter  than  tergal  valves;  dorsal  lobes 
of  cauda  four  in  number,  posterior  pair  very  powerful,  lateral  pair  small,  anterior  median 
pair  lacking. 

Nepionotype.—  Ithaca,  New  York,  March  22,  1913. 

Neanotype. —  With  type. 

Paratypes. —  Numerous  larvae  and  pupae  with  types,  April  26,  1917;  March  22,  1913;  etc. 

Tipula  ignobilis  Loew 

1863     Tipula  ignobilis  Loew.    fieri.  Ent.  Ztschr.,  vol.  7,  p.  280. 

The  adult  flies  of  Tipula  ignobilis  are  not  common  in  collections,  due 
in  part  to  their  retiring  habits.  The  larvae,  however,  are  common  in 
their  preferred  habitat,  saturated  moss  cushions. 

At  Orono,  Maine,  numerous  larvae  were  taken  in  wet  moss  on  June 
17,  1913,  associated  with  other  larvae,  such  as  those  of  Rhaphidolabina, 
Tricyphona,  Pedicia,  and  Tipula  nobilis.  At  Ithaca,  New  York,  on  April 
23,  1917,  four  very  small  larvae  were  found  in  wet  cushions  of  moss  (Ambly- 
stegium).  They  grew  very  rapidly,  emerging  as  adults  on  May  21. 
On  May  22  this  moss  was  carefully  examined  and  about  thirty  fully  grown 
to  rather  immature  larvae  of  this  species  were  taken.  They  were 
associated  with  equally  numerous  larvae  of  Dicranomyia  stulta  O.  S.  At 
Needham's  Glen,  the  species  occurred  in  the  same  moss  that  earlier  in 
the  season  harbored  Tipula  collaris  and  T.  oropezoides.  On  June  14 
two  teneral  adults  of  T.  ignobilis  were  captured,  in  company  with 
Dicranomyia  stulta,  Geranomyia  canadensis,  Dactylolabis  montana,  and 
other  species.  The  species  is  very  common  at  the  Indian  Ladder,  Helder- 
berg  Mountains,  New  York. 

Larva. —  Length,  16.5-18  mm. 
Diameter,  2.5-2.6  mm. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  1011 

Coloration  grayish  brown,  paler  gray  beneath;  thoracic  segments  conspicuously  reddish 
brown. 

Body  terete.  Dorsum  covered  with  a  short,  dark  pubescence,  which  gives  upper  surface 
its  dark  color.  Chaetotaxy  as  follows:  tergites  (Plate  XCVI,  530)  with  a  posterior  trans- 
verse row  of  two  setae  near  base  of  posterior  ring;  sternites  (Plate  XCVI,  531)  with  about 
eight  setae,  anterior  median  row  consisting  of  two  large  setae,  each  with  a  tiny  seta 
proximad  of  it,  the  posterior  pair  of  large  setae  more  widely  separated,  laterad  of  each  of  these 
two  closely  approximated  smaller  setae.  Spiracular  disk  (Plate  XCVI,  532)  surrounded 
by  six  moderately  long  lobes  fringed  with  rather  short,  pale  hairs;  tips  of  ventral  lobes  with 
a  pale  circular  area  provided  with  a  sensory  seta;  inner  face  of  spiracular  disk  with  the 
markings  pale,  ventral  lobes  with  an  indistinct  capillary  brown  line;  two  brown  spots  beneath 
each  spiracle.  Spiracles  large.  Anal  gills  eight,  consisting  of  a  transverse  row  of  four 
short,  slender,  two-branched  lobes  (Plate  XCII,  511). 

Mentum  of  head  capsule  with  median  point  prominent,  the  three  lateral  teeth  of  either 
side  small,  moderately  acute.  Mandible  with  about  four  teeth,  in  addition  to  the  large 
basal  prosthecal  tooth. 

Pupa. —  Length:  male,  12  mm.;  female,  14-15  mm. 

Width,  d.-s.:  male,  1.7-1.8  mm.;  female,  1.8-2  mm. 
Depth,  d.-v.:  male,  1.9-2  mm.;  female,  2.1-2.2  mm. 

Head  and  mouth  parts  as  in  genus.  Pronotal  breathing  horns  rather  long  and  slender, 
tips  a  little  expanded.  Leg  sheaths  rather  short,  just  exceeding  third  abdominal  segment; 
hind  legs  a  little  the  longest,  the  other  tarsi  ending  about  on  a  level. 

Abdominal  tergites  with  armature  weak,  spines  small;  pleurites  and  sternites  with  spines 
notably  larger  and  more  powerful;  sternites  with  sub  terminal  row  consisting  of  twelve  to 
fifteen  spines,  those  of  the  sixth  and  seventh  segments  larger;  no  spines  at  base  of  posterior 
ring;  pleural  spines  setiferous,  very  weak,  one  on  basal  ring,  two  on  posterior  ring.  Male 
cauda  with  ventral  lobes  widely  separated,  terminating  in  slender,  curved  spines  directed 
caudad;  dorsal  lobes  slender,  approximated  on  dorso-median  line.  Female  cauda  (Plate 
XCVI,  533)  with  tergal  valves  a  little  longer  than  sternal  valves,  the  latter  at  their  tips 
terminating  in  slender  points  directed  ventrad  and  laterad;  dorsum  of  cauda  with  the  usual 
six  lobes,  posterior  pair  the  largest,  terminating  in  two  distinct  points;  eighth  segment  on 
pleural  and  sternal  region  with  six  very  powerful,  curved  spines. 

Nepionotype. —  Ithaca,  New  York,  June  3,  1917. 

Neanotype. —  With  larva,  June  3,  1917. 

Paratypes. —  Numerous  larvae  and  pupae,  June  1  to  12,  1917. 

Tipula  abdominalis  (Say) 

1823    Ctenophora  abdominalis  Say.    Journ.  Acad.  Nat.  Sci.  Phila.,  vol.  3,  p.  18. 
Tipula  abdominalis  is  the  largest  eastern  species  of  the  genus,  altho 
some  specimens  of  T.  caloptera  are  nearly  as  large.     The  adult  flies  are 
on  the  wing  in  late  August  and  September,  and  even,  in  fewer  numbers, 
in  June  and  July. 


1012  CHARLES  PAUL  ALEXANDER 

The  large,  fleshy  larvae  occur  beneath  or  among  drift  at  or  near  the 
margins  of  streams.  They  are  usually  abundant  under  saturated  decay- 
ing leaves  or  under  tussocks  of  grass  at  the  edge  of  the  water.  They 
are  entirely  herbivorous,  feeding  on  diatoms,  decaying  plant  tissues, 
and  other  vegetable  matter.  The  larvae  are  readily  distinguished  from 
all  other  species  of  the  genus  by  the  small  spiracles  and  the  bifid  or  split 
-lobes  surrounding  the  spiracular  disk.  Pupation  takes  place  in  the  mud 
or  earth  at  the  water's  edge.  Malloch's  Tipula  sp.  2  (1915-17  b:  200- 
201)  seems  to  refer  to  T.  abdominalis,  but  the  mouth  parts  of  his  speci- 
mens do  not  seem  to  be  normal. 

Larva. —  Length,  55-65  mm. 

Diameter,  7.5-10  mm. 

Coloration  pale  grayish  brown;  anterior  end  darker. 

Form  stout,  terete,  thin-skinned.  Posterior  ring  of  abdominal  segment,  both  above  and 
below,  with  a  naked  transverse  welt,  which  bears  the  setae;  this  welt  very  protuberant  at 
its  lateral  ends,  almost  like  a  pair  of  prolegs.  Skin  naked,  except  for  a  microscopic  dotting 
and  the  rather  short,  delicate  setae.  Chaetotaxy  as  follows:  tergites,  on  welt  at  about  mid- 
length,  a  transverse  row  of  eight  setae,  the  middle  pair  of  each  half  closely  approximated; 
pleurites  with  a  single  seta  opposite  basal  ring,  and  two  opposite  posterior  ring,  arranged 
one  behind  the  other;  sternites  with  four  pairs  of  setae  on  welts,  anterior  middle  pair  closer 
together  than  posterior  lateral  pair.  Spiracular  disk  (Plate  XCVII,  537)  moderately  large, 
flattened,  truncated,  surrounded  by  six  deeply  bifid  and  irregular  lobes;  ventral  lobes  elon- 
gate, with  a  blunt  basal  branch  bearing  two  setae;  posterior  branch  longer,  indistinctly 
bifid  at  tip  and  with  a  few  setae  and  several  hairs;  lateral  lobes  deeply  bifid,  the  ventral 
one  armed  with  lateral  setae;  dorsal  lobes  small,  simple;  inner  faces  of  dorsal  and  lateral 
lobes  with  a  narrow  dark  brown  stripe,  ventral  lobes  with  two  narrow,  usually  indistinct 
lines,  these  markings  broadest  and  darkest  on  dorsal  lobes.  Spiracles  very  small,  circular, 
separated  by  a  distance  about  equal  to  four  times  diameter  of  one.  Anal  gills  six,  long  and 
slender,  the  middle  one  on  either  side  a  little  shorter  than  the  others.  Pleural  region  of 
penultimate  segment  of  body  with  a  blunt  setiferous  tubercle. 

Head  capsule  and  mouth  parts  about  as  in  genus.  Mentum  (Plate  XCVII,  536)  broadly 
transverse,  anterior  margin  with  seven  to  nine  teeth,  in  the  latter  case  the  outermost  teeth 
very  indistinctly  separated  from  the  sublateral  teeth.  Hypopharynx  broad,  anterior  margin 
with  teeth  very  indistinct. 

Pupa. —  Length  of  cast  pupal  skin,  about  35  mm. 

Pronotal  breathing  horns  short,  straight,  cylindrical;  apex  short,  and  but  little  if  at  all 
expanded.  Details  of  mouth  parts  as  in  other  species  of  genus.  Wing  sheaths  ending 
before  apex  of  second  abdominal  segment.  Leg  sheaths  extending  to  about  opposite  end 
of  third  abdominal  segment;  fore  tarsi  the  shortest,  hind  tarsi  the  longest,  ends  of  tarsal 
sheath  thus  forming  a  broad,  inverted  U-shaped  notch. 

Abdominal  armature  generally  weak.  Tergites  (Plate  XCVII,  538)  with  subterminal  row 
broken,  consisting  of  an  anterior  median  pair  of  large  spines,  laterad  of  which  are  two  or 


THE  CRANE-FLIES  OF  NEW  YORK  — PART  II  1013 

three  smaller  spines;  close  to  ends  of  row  a  bifid  setiferous  spine;  pleurites  with  spines  long, 
slender,  directed  strongly  caudad,  tips  narrowly  bifid,  bearing  a  seta  in  notch;  basal  spine 
the  largest;  posterior  spines  two  in  number,  anterior  dorsal  one  the  smaller;  sternites  (Plate 
XCVII,  539)  with  posterior  row  unbroken,  of  comparatively  few  spines,  there  being  about 
eleven  excluding  the  two  larger  spines  at  ends  of  row;  the  innermost  of  the  large  lateral  spines 
conspicuously  bifid,  bearing  a  stout  seta  in  its  notch;  base  of  posterior  ring  on  either  side  of 
median  line  with  a  powerful  conical  spine  which  is  acutely  tipped.  Female  cauda  as  in  genus, 
sternal  valves  conspicuously  shorter  than  the  long  tergal  valves;  the  six  dorsal  lobes  power- 
ful, chitinized,  more  or  less  bifid  at  tips;  dorsal  lateral  lobes  at  end  of  eighth  segment 
split  before  tips. 

Nepionotype. —  Cascadilla  Creek,  Ithaca,  New  York,  May  31,  1913. 
Neanotype. —  Cast  pupal  skin,  reared  at  Ithaca,  September,  1911. 
Paratypes. —  Abundant  larvae  from  type  locality. 

Tipula  taughannock  Alex. 

1915     Tipula  taughannock  Alex.     Proc.  Acad.  Nat.  Sci.  Phila.,  p.  476-479. 

Tipula  taughannock  is  of  exceptional  interest  in  the  striking  color 
dimorphism  that  it  shows,  the  females  being  black  and  yer.ow,  the  males 
light  yellowish.  The  following  account  of  the  habits  of  the  adult  flies 
is  taken  from  the  writer's  field  notes: 

June  12,  1915.  Deciduous  forest  association  of  the  southern  Helderberg  Mountains,  near 
the  village  of  New  Salem,  Albany  County,  New  York.  This  association  is  an  open  deciduous 
forest,  with  an  undergrowth  of  Cystopteris,  Geranium,  Caulophyllum,  and  Impatiens.  It  is  a 
very  open  woods,  having  an  eastern  exposure  and  with  the  talus  slopes  so  old  that  an  extensive 
vegetation  has  sprung  up.  The  great  boulders  scattered  about  thru  the  woods  have  come 
from  the  high  Silurian  and  Devonian  cliffs  above.  The  woods  are  of  such  a  nature  that  much 
sunlight  penetrates  to  the  ground  beneath.  The  forest  cover  shows  a  striking  lack  of  conif- 
erous species,  but  the  following  deciduous  species  are  common:  butternut,  hop  hornbeam, 
hard  maple,  basswood,  white  ash.  The  shrubbery  consists  of  mountain  maple,  bladdernut, 
and  a  few  dogwoods.  The  dominant  herbage  consists  of  jack-in-the-pulpit,  wild  ginger, 
bloodroot,  bishop's-cap,  false  bishop's-cap,  blue  cohosh,  white  baneberry,  herb  robert,  touch- 
me-not,  waterleaf,  bedstraw,  and  other  characteristic  flowering  plants  in  fewer  numbers, 
as  well  as  several  ferns,  such  as  the  bulbous  bladder  fern,  maidenhair,  and,  on  the  rocks, 
the  walking  fern.  The  crane-fly  under  consideration  is  very  common  in  these  woods.  The 
proportion  of  males  to  females  is  about  one  hundred  to  one,  but  this  is  due,  in  large  part 
at  least,  to  the  very  secretive  habits  of  the  latter.  The  males  are  untiring,  almost  always 
moving  along,  silently  and  relentlessly,  in  quest  of  their  mates.  They  pass  in  and  out  among 
the  dense  herbage,  usually  close  to  the  ground,  occasionally  fluttering  up  a  tree  trunk  or  over 
a  mossy  boulder  which  is  covered  with  various  bryophytes  and  walking  ferns.  They  are 
so  intent  upon  their  quest  that  they  are  readily  scooped  up  by  hand.  If  this  is  attempted 
and  fails,  however,  they  become  instantly  alarmed  and  fly  away  with  great  speed,  their  flight 
at  this  time  having  a  strong  undulating  motion.  In  a  position  of  rest,  the  male  almost 
always  hangs  on  the  under  surface  of  a  leaf,  with  the  body  directed  straight  toward  the 
ground.  Several  specimens  of  thia  species,  as  well  as  of  Tipula  trivittata  Say  and  T.  senega 
Alex.,  were  found  dead  in  spider's  webs.  These  small  webs,  made  by  species  of 
Epeiridae  and  Linyphiidae,  are  very  common  on  and  between  the  leaves  of  herbaceous  plants 
and  are  presumably  intended  for  smaller  game.  The  large  Tipulas  are  probably  taken  in 
by  accident. 


1014  CHARLES  PAUL  ALEXANDER 

The  females  are  more  active  when  in  flight  than  are  their  mates  and  are  capable  of  moving 
very  rapidly.  Their  flight  is  a  rapid,  fluttering  progression  along  the  ground.  Copulation 
takes  place  on  either  the  upper  or  the  lower  surface  of  leaves,  usually  near  the  ground.  Some- 
times copulation  is  end  to  end,  with  the  heads  directed  away  from  each  other;  at  other  times 
it  is  face  to  face,  the  bodies  being  arcuated  into  a  convex  loop. 

The  larva  almost  certainly  lives  among  or  beneath  the  decaying  leaves  and  debris  which 
cover  the  talus  slope. 

Tipula  macrolabis  Loew 

1864     Tip-da  macrolabis  Loew.    Berl.  Ent.  Ztschr.,  vol.  8,  p.  58. 

Tipula  macrolabis  is  distinctly  northern  in  its  distribution.  It  is  a 
characteristic  fly  of  northern  deciduous  woods  in  June.  The  following 
notes  were  made  near  the  village  of  Indian  Castle,  Herkimer  County, 
New  York,  on  June  13,  1915: 

A  small  woodland  stream  with  a  forest  cover  of  trees  such  as  hemlock,  beech,  slippery 
elm,  and  basswood,  and  a  ground  cover  of  false  Solomon's  seal,  wood  nettle,  wild  ginger, 
herb  robert,  touch-me-not,  waterleaf,  sarsaparilla,  and  the  two  abundant  ferns  maidenhair 
and  the  bulbous  bladder  fern.  The  males  of  T.  macrolabis  were  in  search  of  the  females, 
and  fluttered  up  the  tree  trunks  often  to  a  height  of  ten  or  fifteen  feet,  flying  close  to  the 
ground,  around  brush  heaps,  hovering  about  the  leafy  ends  of  branches,  and  performing 
similar  actions  in  their  untiring  quest  for  their  mates.  They  occurred  in  company  with 
males  of  T.  fuliginosa  and  T.  valida,  which  were  similarly  engaged  in  searching  for  the  females. 

Tipuline  No.  1  (possibly  Tipula  iroquois  Alex.) 

1863     Tipula  cincta  Loew.     Berl.  Ent.  Ztschr.,  vol.  7,  p.  288-289,  not  T.  cincta  Gmel., 

Syst.  Nat.,  ed.  13,  vol.  1,  p.  2820  (1792). 
1915     Tipula  iroquois  Alex.    Insec.  Inscit.  Menst.,  vol.  3,  p.  128. 

The  larva  discussed  below  has  never  been  reared  and  is  mentioned 
here  principally  because  of  its  interesting  habitat.  It  is  referred  to 
Tipula  iroquois  with  considerable  doubt. 

The  larva  lives  among  dense  mats  of  an  aquatic  moss,  a  Hypnum 
(Rhynchostegium)  of  the  dilatatum  group,  in  the  most  rapid-flowing 
streams.  At  Coy  Glen,  Ithaca,  New  York,  these  larvae  are  especially 
frequent,  often  living  at  the  brink  of  falls  or  rapids  in  the  most  rushing 
waters.  A  study  of  the  structure  of  the  larva  reveals  numerous  small 
but  prominent  tubercles,  which  doubtless  assist  the  insect  in  clingi-ng  to 
the  moss  stems.  The  gills  are  large,  but  no  better  developed  than  in 
many  aquatic  species  of  the  genus  that  live  in  much  less  lotic  conditions. 
The  green  color  and  the  transverse  rows  of  tubercles  on  the  body  give  the 
larva  a  strong  resemblance  to  its  mossy  habitat.  The  larvae  are  very 
sluggish  and  crawl  but  slowly,  often  appearing  quite  dead  for  long  periods 
of  time. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  1015 

Associated  with  these  larvae  in  Coy  Glen,  the  following  characteristic 
forms  of  insect  life  occur : 

Plecoptera:  a  small  species  of  Perlidae. 

Ephemerida:  Ephemeridae,  such  asBaetis,/ron/m^7zs,  Ephemerella, 

and  others. 

Trichoptera:  Ryacophilidae,  Hydroptilidae,  and  other  forms. 
Neuroptera:  Chauliodes  larvae. 

Diptera:    Chironomidae,    a   few;   Psychodidae,    Psychoda   alhitarsis 
Banks;  Stratiomyiidae;  Anthomyiidae,  Limnophora  torreyae  Job.; 
and  other  groups. 
Coleoptera:  Parnidae,  larvae  of  Psephenus  lecontei  (Lee.),  and  adult 

beetles  of  a  species  of  Elmis  in  large  numbers. 

Up  to  the  present  time  it  has  been  found  impossible  to  rear  this  larva 
to  the  adult  condition,  chiefly  because  of  the  constant  need  of  well-aerated 
water  and  the  difficulty  of  supplying  it.  By  placing  the  larvae  in  the  folds 
of  moistened  cheesecloth,  it  was  possible  on  one  occasion  to  carry  the 
species  to  the  pupal  state,  but  no  further.  It  will  be  of  interest  to 
ascertain  the  identity  of  this  conspicuous  larva. 

Larva. —  Length,  24-25  mm. 
Diameter,  2.6-3  mm. 

Coloration  above,  dark  green  with  a  brown  pattern;  beneath,  light  green  with  indistinct 
transverse  brown  lines;  on  dorsum  a  pale  longitudinal  mark  on  sides  of  posterior  ring,  cross- 
ing sutures  between  segments  onto  extreme  base  of  anterior  ring  of  following  segment,  the 
dark  area  of  each  segment  thus  appearing  cruciform,  this  cross-shaped  mark  spotted  and 
marbled  with  darker  in  transverse  rows;  base  of  gills  and  center  of  spiracular  disk  light  green 
in  living,  healthy  larvae. 

Form  moderately  terete,  each  segment  with  transverse  rows  of  small,  prominent  tubercles, 
some  of  which  are  provided  with  setae.  On  dorsum  of  posterior  ring  a  subterminal  row  of 
six  tubercles,  the  middle  one  on  each  side  with  two  setae,  the  remaining  four  tubercles  unise- 
tose;  two  rows  of  smaller  naked  tubercles  at  base  and  middle  of  posterior  annulus;  basal 
annulus  with  four  transverse  rows  of  small  naked  tubercles;  pleurites  with  three  tubercles, 
a  small  seta  on  basal  ring  ventrad  of  basal  tubercle,  and  two  setae  on  posterior  ring  ventrad 
and  cephalad  of  posterior  tubercle;  sternites  on  posterior  ring  with  six  setae,  four  on  ante- 
rior row,  the  two  middle  ones  very  tiny.  Spiracular  disk  almost  as  in  T.  collaris,  T.  ignobilis, 
and  similar  species,  surrounded  by  six  lobes  fringed  with  moderately  long  hairs;  inner  face 
of  lobes  somewhat  pale;  ventral  lobes  with  a  narrow,  capillary,  dark  brown  line,  extending 
from  tip  toward  base;  lobes  narrowly  and  more  or  less  indistinctly  margined  with  brown; 
two  brown  spots  at  base  of  each  ventral  lobe,  underneath  each  spiracle;  dorsal  and  lateral 
lobes  jutting  backward  at  tips  into  fleshy  conical  points.  Spiracles  circular,  moderately 
large,  separated  by  a  distance  about  equal  to  one  and  one-half  diameter  of  one.  Anal  gills 
with  eight  branches,  rather  short  and  stout,  with  two  lateral  divergent  branches  on  either 
side  and  an  inner  pair  with  one  ventral  and  one  posterior  branch  (Plate  XCII,  512). 


1016  CHARLES  PAUL  ALEXANDER 

Head  capsule  and  mouth  parts  of  almost  normal  tipuline  appearance.  Mentum  with 
outer  plate  forming  the  unusually  long  apical  point,  inner  plate  adding  three  teeth  on  either 
side,  margins  bulging.  Antenna  with  apical  papillae  somewhat  flattened,  surrounded  by 
three  sense  pegs. 

(Described  from  abundant  specimens,  Coy  Glen,  Ithaca,  New  York,  April  23,  1914.) 

Tipuline  No.  2 

The  larva  discussed  below  is  known  only  from  a  single,  apparently 
fully  grown,  specimen.  The  writer  has  no  clue  as  to  which  species  it 
represents,  altho  from  the  larval  structure  it  is  obviously  allied  to  Tipula 
selene  Meig.  of  Europe. 

Larva. —  Length,  23  mm. 
Diameter,  3  mm. 

Coloration,  a  rather  uniform  pale  yellowish  or  reddish  brown. 

Body  covered  with  a  rather  sparse,  long  pubescence,  setae  unusually  long  and  powerful. 
Chaetotaxy  as  follows:  tergites  (Plate  XCVI,  534)  with  two  strong  lateral  setae  near  pos- 
terior margin  and  an  additional  one  at  extreme  lateral  margin  of  ring;  pleural  setae,  one  on 
basal  ring,  two,  one  behind  the  other,  on  posterior  ring;  sternites  with  a  transverse  pair  of 
powerful  setae  near  extreme  lateral  margin,  and  two  anterior  pairs  of  much  smaller  setae. 
Spiracular  disk  (Plate  XCVI,  535)  with  four  elongated,  cylindrical,  chitinized  horns  which 
are  narrowed  to  the  acute,  blackened,  slightly  curved  tips;  longer  dorsal  pair  rather  closely 
approximated,  lying  almost  parallel,  with  tips  a  little  curved  dorsad;  shorter  and  more 
slender  lateral  horns  directed  ventrad  at  tips,  at  base  on  inner  face  with  a  powerful  seta, 
the  large  black  spiracles  lying  above  base  of  lateral  horns;  ventral  lobes,  if  present,  very 
blunt  and  indistinct.  Anal  gills  not  protruded  in  the  only  specimen  available. 

Head  capsule  and  mouth  parts  rather  normal  but  showing  the  following  points  of  difference 
from  the  usual  Tipula  type:  Mentum  almost  completely  split,  apical  point  long  and  narrow; 
the  three  lateral  teeth  on  either  side  blunt,  lateral  pair  tending  to  be  reduced. 

Hypopharynx  with  five  blunt  teeth.  Antenna  short,  stout,  length  only  about  twice 
diameter,  at  tip  with  a  subglobular,  feebly  chitinized  papilla. 

(Described  from  a  single  larva  found  beneath  a  stone  in  a  field  near  Taughannock  Falls, 
Tompkins  County,  New  York,  May  1,  1912.) 

Genus  Nephrotoma  Meigen  (Gr.   kidney  +  I  cut) 

1800     Pales  Meig.     Nouv.  Class.  Mouch.,  p.  14  (nomen  nudum). 

1803     Nephrotoma  Meig.     Illiger's  Mag.,  p.  262. 

1834     Pachyrrhina  Macq.     Hist.  Nat.  Ins.,  Dipt.,  vol.  1,  p.  88. 

The  large  genus  Nephrotoma,  including  some  one  hundred  and  fifty 
described  species,  is  very  close  to  Tipula  in  all  respects.  The  writer 
cannot  attempt  to  separate  the  immature  stages  of  the  genus  from  those 
of  Tipula,  on  the  scanty  material  that  has  been*  available  for  study. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  1017 

In  Europe,  Beling  and  others  have  described  the  life  histories  of  about 
eight  of  the  commoner  species.  The  following  species  live  in.  earth, 
beneath  a  covering  of  leaf  mold:  analis  (Schum.),  cornitina  (Linn.),  lineata 
(Scop.),  maculata  (Meig.),  lunulicornis  (Schum.),  pratensis  (Linn.).  The 
following  have  been  described  as  living  in  decaying  wood,  some  of  the 
records  apparently  being  in  error:  cornicina  (Linn.),  crocata  (Linn.), 
lineata  (Scop.),  quadrifaria  (Meig.).  Some  of  the  species  are  injurious 
to  young  seedlings,  especially  those  of  coniferous  plants. 

In  North  America  the  commonest  species,  N.  ferruginea  (Fabr.),  has 
been  discussed  several  times  in  its  economic  relations  to  agriculture.  Hart 
(1898  [1895] :  218-219)  gives  an  excellent  description  of  the  immature 
stages,  while  Malloch  (1915-17  b:  206)  gives  supplementary  notes  and 
figures  of  the  same  species.  The  immature  stages  live  in  sand  or  earth. 
N.  virescens  (Loew)  was  bred  from  a  larva  found  in  moss  OB.  Plummers 
Island,  Maryland,  on  April  5,  1913,  by  R.  C.  Shannon.  N.  eucera  (Loew) 
and  N.  polymera  (Loew)  have  been  reared  from  larvae  taken  under  leaf 
mold  in  woods  by  Mabel  M.  Alexander. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  1019 


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VERRALL,  G.  H.  Another  hundred  new  British  species  of  Diptera« 
Ent.  mo.  mag.  48:20-27.  (Reference  on  p.  23.)  1912. 

VIALLANES,  H.  Sur  1'appareil  respiratoire  et  circulatoire  de  quelques 
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1880. 

VILLENEUVE,  J.  Anomalie  analogue  a  la  j>recedente  constatee  chez  UE 
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ent.  Ztg.  29:81-82.  1910. 

VIMMER,  A.  Pachyrhina  iridicolor  Schummel  jako  skudce  fepy  cukrove 
Casopis  Ceske  Spolecnosti  Ent.  2:70-75.  1905. 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II  1041 

— SrovnaVaci  studie  o  ustnim  ustroji  larev  Pachyrin  a  Tipulin. 

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WAHL,  B.     Schnakenlarven    als    Pflanzenschadlinge.     K.    Pflanzensch, 
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WAHLGREN,  EINAR.     Det  olandska  alvarets  djurvarld.     Ark.  Zool.  II1: 
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12 


1042  CHARLES  PAUL  ALEXANDER 

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ZETTERSTEDT,  JOHANN  WILHELM.  Diptera  Scandinaviae  disposita  et 
descripta  10:3711^090.  (Reference  on  p.  4014,  4016,  4023.)  1851. 

EXPLANATION  OF  ABBREVIATIONS  USED  ON  PLATES 

A = antenna  PI  =  pleuron 

C = cardo  PM  =  prementum 

Lb=labium,  or  labial  lobes  S  =  stipes 

M  =  men  turn  SM  =  submentum 

Md  =  mandible  Sp  =  spiracle 

MX  =  maxilla  St  =  stern  ite 

P  =  palpus,  or  palpal  sheath  T  =  tergite 


Memoir  34,  An  Economic  Study  of  Farm  Layout,  the  fourth  preceding  number  in  this  series  of  publica- 
tions, was  mailed  on  January  31,  1921. 

Memoir  35,  Some  Effects  of  Potassium  Salts  on  Soils,  the  third  preceding  number  in  this  series  of  publica- 
tions, was  mailed  on  January  29,  1921. 

Memoir  36,  Resistance  of  the  Roots  of  Some  Fruit  Species  to  Low  Temperature,  the  second  preceding  number 
in  this  series  of  publications,  was  mailed  on  January  19,  1921. 

Memoir  37,  A  Modified  Babcock  Method  for  Determining  Fat  in  Butter,  the  next  preceding  number  in  this 
series  of  publications,  was  mailed  on  December  10,  1920. 


MEMOIR  38 


PLATE  XII 


rO/77C7/-/<7 


4r? 


1,  Eriocera  spinosa  emerging  from  pupal  hull. 

2-3,  Spiracular  disk  and  tracheation  of  larvae:     2,  Antacha 
saxicola  (apneustic);   3,   Dicranota   Umaculata   (metapneustic), 
after  Miall 

4,  Phylogenetic  tree 

1013 


MEMOIR  38 


PROTOPLASA    FITCHII,    SUPPOSITION 

5,  Dorsal   aspect;  6,   spiracular  disk;  7,  head,  dorsal  aspect;  8,  head,  ventral  aspect; 

anal  gill 

1044 


PLATE  XIV 


10 


12 


15 


PTYCHOPTERA    RUFOCINCTA 


Larva:    10,  dorsal  aspect;  11,  breathing  tube  extended;  12,  head,  ventral  aspect 
Pupa:     13,  lateral  aspect;  14,  ventral  aspect;  15,  female  cauda,  ventral  aspect 

1045 


1046 


1047 


n 

1 


I      ° 
I      O 

- 


1048 


MEMOIR  38 


PLATE  XVIII 


46 


47 


BITTACOMORPHA    CLAVIPES 


43 


Larva:    41,  labium;  42,  mandible  and  antenna,  dorsal  aspect 

Pupa:    43,  ventral  aspect;  44-46,  types  of  abdominal  tubercles;  47,  arrangement  of  leg 
sheaths;  48,  male  cauda,  dorsal  aspect 


1049 


MEMOIR  38 


PLATE  XIX 


49 


56 


TBICHOCEBA    SP.    (SEGELATIONIS,    SUPPOSITION) 

Larva:     49,  dorsal  aspect;  50,  spiracular  disk,  lateral  aspect;  51,  spiracular  disk,  dorsal 

aspect;  52,  head,  ventral  aspect  (after  De  Meijere) ;  53,  head,  dorsal  aspect  (after  De  Meijere) 

Pupa:    54,  lateral  aspect;  55,  female,  ventral  aspect;  56,  female  cauda,  lateral  aspect 

1050 


MEMOIR  38 


PLATE  XX 


ANTOCHA    8AXICOLA 


Larva:     57,  dorsal  aspect;  58,  head  capsule,  dorsal  aspect;  59,  mentum 
Pupa:     60,  ventral  aspect;  61,  pronotal  breathing  horn,  lateral  aspect;  62,  female  cauda, 
lateral  aspect 


1051 


1052 


MEMOIR  38 


PLATE  XXII 


69 


71 


70 


ANTOCHA   SAXICOLA,    PUPA 

),  Lateral  aspect;    70,  male  cauda,  dorsal  aspect;    71,  fifth  abdominal  segment,  dorsal 

aspect  (diagrammatic) 


'1053 


CO 


o 
•a 

ll 

s  a 
«  § 


§  1- 


1054 


MEMOIR  38 


PLATE  XXIV 


80 


83 


LIMNOBIA    CINCTIPES 


Larva:     79,  head  capsule,  dorsal  aspect;  80,  hypopharynx;  81,  mentum;  82,  antenna;  83, 
spiracular  disk 
Pupa:    84,  female,  lateral  aspect;  85,  female  cauda,  dorsal  aspect 


1055 


MEMOIR  38 


PLATE  XXV 


88 


LIMNOBIA    CINCTIPES 


Larva:  86,  labrum-epipharynx;  87,  mandible  and  maxilla;  88,  mandible;  93,  spiracular 
disk,  dorsal  aspect 

Pupa:  89,  female,  ventral  aspect;  90,  pronotal  breathing  horn,  lateral  aspect;  91,  male 
cauda,  lateral  aspect;  92,  male  cauda,  dorsal  aspect 


1056 


MEMOIR  38 


PLATE  XXVI 


94 


100 


101 


102 


LIMNOBIA   TRIOCELLATA   AND    L.    FALLAX 


Limnobia  triocellata:     94,   spiracular  disk 

Limnobia  fallax,  larva:  95,  labrum-epipharynx;  96,  mentum;  97,  antenna;  98,  mandible; 
99,  maxilla 

Limnobia  fallax,  pupa:  100,  mouth  parts;  101,  pronotal  breathing  horn;  102,  female 
cauda,  dorsal  aspect;  103,  female  cauda,  lateral  aspect 

1057 


1058 


1059 


MEMOIR  38 


PLATE  XXIX 


120 


DICBANOMYIA     BADIA,    PUPA 

118,  Lateral  aspect;  119,  mouth  parts;  120,  female  cauda,  dorsal  aspect 


1060 


1061 


MEMOIR  38 


PLATE  XXXI 


128 


136 


RHIPIDIA  BRTANTI,  RHAMPHIDIA  MAINENSIS,   AND   RHAMPHIDIA   FLAVIPES 

Rhipidia  bryanti,  pupa:  128,  mouth  parts;  129,  pronotal  breathing 
horn;  130,  male  cauda,  lateral  aspect;  131,  male  cauda,  dorsal  aspect 

Rhamphidia  mainensis,  larva:  132,  lateral  aspect;  133,  inentum; 
135,  mandible;  136,  spiracular  disk 

Rhamphidia  flavipes,  larva:     134,  antenna 

1062 


1063 


MEMOIR  38 


PLATE  XXXIII 


143 


144 


RHAMPHIDIA   FLAVIPES,    PUPA 

142,  Lateral  aspect;  143,  fifth  abdominal  segment,  dorsal  aspect  (diagrammatic);  144, 
fifth  abdominal  segment,  ventral    aspect  (diagrammatic) 


1064 


MEMOIR  38 


PLATE  XXXIV 


153 


ULA   ELEGANS,    LARVA 

145,  Lateral  aspect;  146,  head  capsule,  dorsal  aspect;  147,  labrum-epipharynx;  148,  mentum 
149,  antenna;  150,  mandible,  lateral  aspect;  151,  mandible,  from  inside;  152,  maxilla;  153, 
spiracular  disk 


1055 


1066 


PLATE  XXXVI 


160 


161 


159 


EPIPHBAGMA   SOLATRIX,    LARVA 


158,  Head  capsule,  ventral  aspect;  159,  mentum;    160,  prementum;  161,  hypopharynx; 
162,  maxilla;  163,  spiracular  disk 


1067 


MEMOIR  38 


PLATE  XXXVII 


166 


PI 


Sp 


167 


EPIPHRAGMA    SOLATRIX   'AND    E.    FASCIPENNIS 


Epiphragma  solatrix,  pupa:     164,  lateral  aspect;  166,  head  of  male,  ventral  aspect;  167, 
fifth  abdominal  segment  (diagrammatic) 

Epiphragma  fascipennis,  pupa:      165,  cephalic  crest,  lateral  aspect 

1068 


1069 


MEMOIR  38 


PLATE  XXXIX 


PSEUDOLIMNOPHILA    LUTEIPENNIS    AND    P.    INORNATA 

Pseudolimnophila  luteipennis,  larva:     172,  mentum;  173,  hypopharynx;  175,  mandible; 
176,  spiracular  disk 

Pseudolimnophila  inornata,  larva:     174,  antenna 

1070 


1071 


MEMOIB  38 


PLATE  XLI 


185 


184 


186 


187 


188 


DACTYLOLABIS    DENTICULATA,    D.    WODZICKII,    AND    D.    CUBITALIS 

Dactylolabis  denticulate  (after  Mik):  181,  pupa;  182,  larva,  head  capsule,  ventral  aspect; 
183,  larva,  dorsal  aspect 

Dactylolabis  wodzickii  (after  Nowicki):  184,  pupa;  185,  larva,  head  capsule;  186,  larva, 
spiracular  disk 

Dactylolabis  cubitalis,  pupa:  187,  pronotal  breathing  horn;  188,  second  abdominal  seg- 
ment, showing  spiracle;  189,  male  cauda,  dorsal  aspect 

1072 


MEMOIR  38 


PLATE  XLII 


LIMNOPHILA  (DICRANOPHEAGMA)  FUSCOVARIA,  LARVA 

190,  Cephalic  and  caudal  ends,  dorsal  aspect;  191,  head  capsule,  ventral  aspect;  192,  labrum 
epipharynx;  193,  antenna;  194,  mandible;  195,  spiracular  disk 


1073 


13 


3 


V  ' 

3i 
>^- 

•T 

•x*' 

-*— 

\, 

>^ 

§8  ^ 

a  ^ 

>j* 

2  ^r 

H    ^                                          ^ 

3-^-  -^ 

1074 


1075 


MEMOIR  38 


PLATE  XLV 


LIMNOPHILA  (LASIOMASTIX)  MACROCERA,  PUPA 

>. 

209,  Male,  lateral  aspect;  210,  mouth  parts;  211,  male  cauda,  dorsal  aspect;  212,  female 

cauda,  lateral  aspect 


1076 


MEMOIR  38 


PLATE  XLVI 


V        V 


213 


214 


215 


217 

LIMNOPHILA   PICTIPENNIS,    L.    PUNCTATA,   AND   ULOMORPHA    PILOSELLA 


219 


Limnophila  pictipennis:    213,  head  capsule  (after  Brauer) 
Limnophila  punctata:     214,  spiracular  disk  (after  Gerbig) 

Ulomorpha  pilosella,  larva:     215,  head  capsule,  dorsal  aspect;  216,  labrum;  217,  antenna; 
218,  mandible;  219,  spiracular  disk,  dorsal  aspect 

1077 


1078 


1079 


MEMOIR  38 


PLATE  XLIX 


233 


234 


235 


236 


PILARIA    TENUIPES,    PUPA 


233,  Female,  lateral  aspect;  234,  female  cauda,  lateral  aspect;  235,  fifth  abdominal  segment, 
dorsal  aspect;  236,  female  cauda,  dorsal  aspect 


1080 


s 

CM 


0     0  0* 


0 


?*  s 


#:\ 


0 


sf 


:l- 
!1 

!S 


If 


1081 


MEMOIR  38 


243 


PLATE  LI 


245 


248 


244 


246 


249 


HEXATOMA   MEGACERA 


247 


Larva:     243,  labrum;  244,  antenna;  245,  mandible;  246,  spiracular  disk 

Pupa:    247,  lateral  aspect;  248,  male,  ventral  aspect;  249,  female,  ventral  aspect 


1082 


5  i 


1 

S  -/  i 
252 

HEXATOMA  MEGACERA  AND  ERIO 

Hexatoma  megacera:  250,  larva,  apex  of  labrum;  251,  Ian 
segment,  lateral  aspect 
Eriocera  cinerea,  larva:  253,  labrum;  254,  pharyngeal  pi 

fc 

—  ^ 

^ 

^      1 

-* 
^ 

£                         ^" 
"**              V- 

- 

*—                ^ 
i* 

L   f 

1083 


MEMOIR  38 


PLATE  LIII 


260 


261 


ERIOCERA    CINEREA,    PUPA 

256,  Male,  lateral  aspect;  257,  cephalic  crest  of  male,  ventral  aspect;  258,  mouth  parts; 
259,  fifth  abdominal  segment,  lateral  aspect;  260,  male  cauda,  dorsal  aspect;  261,  female 
cauda,  dorsal  aspect 

1084 


MEMOIR  38 


PLATE  LIV 


267 


264 


268 


269       270 


271 


ERIOCERA    SPINOSA,    E.    CINEREA,    E.    LONGICORNIS,    AND    E.    FULTONENSIS 

Eriocera  spinosa,   larva:     262,   spiracular  disk;  207,   head   capsule,  dorsal  aspect;  271, 
nandible 

Eriocera  cinerei,  larva:     263  and  234,  spiracular  disk;  270,  mandible 
Eriocera  longicornis,  larva:     285,  spiracular  disk 
Eriocera  fultonensis,  larva:     236,  spiracular  disk;  268,  labrum;  269,  mandible 


1085 


MEMOIR  38 


PLATE  LV 


278 


277 


272 


273 


279 


276 


282 


280 


ERIOCERA   LONGICORNIS,    E.    FULTONENSI3,    AND    E.    SPINOSA 


Eriocera  longicornis,  pupa:  272,  male,  lateral  aspect;  273,  male,  ventral  aspect;  274, 
female  cauda,  lateral  aspect;  275,  head  of  male,  ventral  aspect;  277,  thorax  of  male,  dorsal 
aspect;  278,  female,  lateral  aspect 

Eriocera  fultonensis,  pupa:     276,  female  cauda,  lateral  aspect;  279,  female,  lateral  aspect 
Eriocera  spinosa,  pupa:     280,  female,  lateral  aspect;  281,  male  cauda,  dorsal  aspect;  282, 
male  cauda,  ventral  aspect 


1086 


MEMOIR  38 


PLATE  LVI 


PI 


1 

1             tj 

t     /SP 

/I 

^  . 

V'Wj/M  yi/\/iAfA 

l  ^"""""-""i 

285 


5t 


ft 


288 


287 


ERIOCERA    SPINOSA,    E.    LONGICORNIS,    AND   E.    FULTONENSIS 

Eriocera  spinosa,  pupa:  283,  fifth  abdominal  segment,  lateral  aspect  (diagrammatic); 
284,  female  cauda,  dorsal  aspect 

Eriocera  longicornis,  pupa:  285,  fifth  abdominal  segment,  lateral  aspect  (diagrammatic); 
283,  male  cauda,  lateral  aspect 

Eriocemfultonensis,  pupa :  287,  cephalic  crest  of  male,  ventral  aspect;  288,  fifth  abdominal 
segment,  lateral  aspect  (diagrammatic) 

1087 


MEMOIR  38 


PLATE  LVII 


289 


292 


294 


295 


293 


PENTHOPTEBA    ALBITARSIS 


Larva:     289,  labrum;  290,  antenna;  291,  mandible;  292,  spiracular  disk,  dorsal  aspect; 
293,  spiracular  disk,  lateral  aspect 

Pupa:    294,  pronotal  breathing  horn;  295,  male  cauda,  lateral  aspect 


1088 


MEMOIR  38 


PLATE  LVIII 


299 


PENTHOPTERA   ALBITARSIS,    PUPA 

296,  Female,  lateral  aspect;  297,  female,  ventral  aspect;  298,  female  cauda,  dorsal  aspect; 
299,  male  cauda,  lateral  aspect 


1089 


MEMOIR  38 


PLATE  I.IX 


304 


303 


302 


300 


305 


ADELPHOMYIA    MINUTA    (SUPPOSITION),    LARVA 

300,  Labrum;  301,  antenna;  302,  mentum;  303,  mandible;  304,  maxilla;  305,  spiracular  disk 


1090 


MEMOIB  38 


PLATE  LX 


307 


.310 

^^^^^^  ADELPHOMYIA   MINUTA    (SUPPOSITION),  PUPA 

306,  Female,  lateral  aspect;  307,  mouth  parts;  308,  female  cauda,  dorsal  aspect;  309,  male 
cauda,  dorsal  aspect;  310,  male  cauda,  lateral  aspect 


1091 


MEMOIR  38 


PLATE  LXI 


311 


315 


PEDICIA   ALBIVITTA,    LARVA 


312 


311,  Dorsal  aspect;  312,  head  capsule,  ventral  aspect;  313,  antenna;  314,  maxillary  palpus; 

315,  anal  gills,  ventral  aspect 


1092 


MEMOIR  38 


PLATE  LXII 


324 


325 


322 


EHAPHIDOLABINA   FLAVEOLA 


Larva:  316,  mentum;  317,  hypopharynx;  318,  antenna;  319,  mandible;  320,  maxilla; 
321,  spiracular  disk 

Pupa:  322,  mouth  parts;  323,  pronotal  breathing  horn,  dorsal  aspect;  324,  pronota! 
breathing  horn,  lateral  aspect;  325,  male  cauda,  dorsal  aspect 


1093 


MEMOIR  38 


PLATE  LXII 


334 


TRICYPHONA  INCONSTANS 


Larva:  326,  mentum;  327,  hypopharynx;  328,  antenna;  329,  mandible  and  maxilla: 
330,  spiracular  disk 

Pupa:  331,  mouth  parts;  332,  pronotal  breathing  horn,  dorsal  aspect;  333,  pronotai 
breathing  horn,  lateral  aspect;  334,  female  cauda,  lateral  aspect 

1094 


M 


M  — 

O    3  S5 


II 


ill 


rO 


. 

III 

IB 

a  * 


g*e* 

co  ^-^ 

111 


1095 


MEMOIR  38 


PLATE  LXV 


344 


345 


349 


346 

MOLOPHILUS   HIBTIPENNIS,    LARVA 

344,  Lateral  aspect;  345,  head  capsule,  ventral  aspect;  346,  mental  plate;  347,  hypopharynx; 
348,  antenna;  349,  mandible;  350,  maxilla;  351,  spiracular  disk 


1096 


MEMOIR  38 


PLATE  LXVI 


355 


352 


354 


356 


MOLOPHILUS   HIRTIPENNIS,    PUPA 

352,  Female,  lateral  aspect;  353,  male  cauda,  lateral  aspect;  354,  male  cauda,  dorsal  aspect; 
355,  female  cauda,  lateral  aspect;  356,  female  cauda,  dorsal  aspect 


1097 


MEMOIR  38 


PLATE  LXVII 


357 


360 


359 


T                PI 

5r 

•     • 

^Wfr/w^A/'rrr*'*""  ^ 

J&               * 

,,_ 

358 


361 


362 


363 


ERIOPTERA    MEGOPHTHALMA 


Larva:    357,  spiracular  disk;  358,  anal  gills,  ventral  aspect 

Pupa:  359,  female,  lateral  aspect;  360,  female,  ventral  aspect;  361,  fifth  abdominal  seg- 
ment, lateral  aspect  (diagrammatic);  362,  male  cauda,  lateral  aspect;  363,  male  cauda,  dorsal 
aspect 

1098 


1099 


MEMOIR  38 


PLATE  LXIX 


369 


373 


370 


ERIOPTERA    CHLOROPHYLLA,    E.    SEPTEMTRIONIS,    AND   E.    ARMATA 

Erioptera  chlorophylla,  pupa:     369,  female,  lateral  aspect;  370,  female  cauda,  lateral  aspect 
Erioptera  septemtrionis,  pupa:    371,  male  cauda,  lateral  aspect;  372,  male  cauda,  dorsal 
aspect 

Erioptera  armata,  pupa:     373,  female,  lateral  aspect 

1100 


MEMOIR  38 


PLATE  LXX 


376 


374 


378 


377 

ORMOSIA   NUBILA,    O.    INNOCENS,    AND    O.    MEIQENII 

Ormosia  nubila,  larva:     374,  mandible;  375,  spiracular  disk 

Ormosia   innocens,   pupa:    376,    female,    lateral   aspect;   377,    male  cauda,   dorsal 
aspect;  378,  male  cauda,  lateral  aspect 

Ormosia  meigenii,  larva:     379,  spiracular  disk 

1101 


MEMOIR  38 


PLATE  LXXI 


380 


381 


382 


ORMOSIA    NUBILA,    PUPA 

380,  Female,  lateral  aspect;  381,  cephalic  crest  of  female,  ventral  aspect;  382,  female 

cauda,  dorsal  aspect 

1102 


MEMOIR  38 


PLATE  LXXII 


385 


386 


384 


387 


ORMOSIA   NIGRIPILA,    PUPA 

383,  Male,  lateral  aspect;  384,  pronotal  breathing  horn;  385,  fifth  abdominal  segment, 
lateral  aspect  (diagrammatic);  386,  male  cauda,  dorsal  aspect;  387,  female  cauda,  lateral 
aspect 


1103 


MEMOIR  38 


PLATE  LXXIII 


395 


*'      VI 

\ 

f  rr             N       1 

r1     Nd 

\ 

f 

391 


394 


HELOBIA   HYBRIDA   AND    TRIMICRA    PILIPES 


HeloUa  hybrida,  larva:    388,  spiracular  disk 

Helobia  hybrida,  pupa:  389,  female,  lateral  aspect;  390,  mouth  parts;  391,  arrangement 
of  leg  sheaths;  392,  male  cauda,  dorsal  aspect;  393,  female  cauda,  lateral  aspect;  394,  male 
cauda,  lateral  aspect 

Trimicra  pUipes,  larva:    395,  spiracular  disk  (after  Gerbig) 


1104 


1105 


14 


MEMOIR  38 


PLATE  LXXV 


403 


402 


404 


405 


406 


GONOMTIA   ALBXANDERI    AND    ERIOPTERINE    NO.    1 


Gonomyia  alexanderi,  larva:    402,  mandible;  403,  spiracular  disk 
Eriopterine  No.  1,  larva:    404,  lateral  aspect;  405,  spiracular  disk  and  anal  gills; 
406,  lobe  of  spiracular  disk,  enlarged 


1106 


1107 


II 


P.  be 

II 


• 


52 

$rh 


P 
- 


1108 


MEMOIR  158 


PLATE  LXXVIII 


420 


421 


422 


423 


TRENTEPOHLIA    PENNIPES   AND    T.    BROMELIADICOLA 


Trentepohlia  pennipes,  pupa:  419,  lateral  aspect  (after  De  Meijere) 
Trentepohlia  bromeliadicola,  larva  (after  Picado) :  420,  anal  gills 

Trentepohlia  bromeliadicola,  pupa  (after  Picado):  421,  female,  ventral  aspect;  422,  male 
cauda,  lateral  aspect;  423,  female  cauda,  lateral  aspect 


1109 


1110 


1111 


MEMOIR  38 


PLATE  LXXXI 


442 


ELEPHANTOMYIA    WESTWOODI,    PUPA 

441,  Female,  lateral  aspect;  442,  female,  ventral  aspect 


1112 


MEMOIR  38 


PLATE  LXXXII 


444 


445 


446 


447 


443 


CHIONEA   ARANEOIDES,    LARVA    (AFTER   BRAUER) 

443,  Dorsal  aspect;  444,  mandible;  445,  possibly  mental  plate;  446,  spiracular  disk, 
lateral  aspect;  447,  spiracular  disk,  dorsal  aspect 

1113 


MEMOIR  38 


PLATE  LXXXIII 


452 


449 


450 


PHALACROCEBA    REPLICATA    AND    TRIOGMA    TRISULCATA 


Phalacrocera  replicata,  larva:  448,  lateral  aspect 

Phalacrocera  replicata,  pupa:  449,  male,  lateral  aspect;    450,  male,  dorsal  aspect;  451, 
mouth  parts 

Triogma  trisulcata:  452,  larva  (after  Steinmann) 

1114 


MEMOIR  38 


PLATE  LXXXIV 


453 


454 


456 


CYLINDROTOMA    SPLENDENS 


455 


Larva:  453,  dorsal  aspect;  454,  mandible 

Pupa:  455,  female,  lateral  aspect;  456,  head  of  female,  ventral  aspect 


1115 


1116 


1117 


MEMOIR  38 


PLATE  LXXXVII 


474 


475 


477 


BRACHTPBEMNA    DISPELLENS,    PUPA 

474,  Head,  ventral  aspect;  475,  pronotal  breathing  horn;  476,  arrangement  of  leg  sheaths; 
477,  male  cauda,  ventral  aspect 


1118 


MEMOIR  38 


PLATE  LXXXVIII 


484 


482 


TANYPTEBA    FRONTALIS 

Larva:  478,  mentum;  479,  antenna;  480,  mandible;  481,  fifth  abdominal  segment,  dorsal 
aspect  (diagrammatic) ;  482,  fifth  abdominal  segment,  ventral  aspect  (diagrammatic) ;  483, 
spiracular  disk 

Pupa:  484, "female,  lateral  aspect;  485,  mouth  parts;  486,  pronotal  breathing  horn;  487, 
female  cauda,  dorsal  aspect 

1119 


MEMOIR  38 


PLATE  LXXXIX 


492 


488 


491 


LONGURIO    TESTACEUS    AND    AESHNASOMA    RIVERTONENSIS 

Longurio  testaceus,  larva:  488,  mentum;  489,  hypopharynx;  490,  spiracular  disk 
Longurio  testaceus,  pupa:  491,  male,  lateral  aspect 

Aeshnasoma  rivertonensis,  larva:  492,  spiracular  disk,  showing  branched  anal  gills  (after 
Johnson) 


1120 


MEMOIR  38 


PLATE  XC 


496 


497 


495 


493 


494 


HOLORUSIA   RUBIGINOSA, 


Larva:  493,  men  turn;  494,  hypopharynx;  495,  antenna;  496,  spiracular  disk 
Pupa:  497,  male,  lateral  aspect 

1121 


MEMOIR  38 


498 


505 


PLATE  XCI 

504 


PRIONOCERA    FUSCIPENNIS 

Larva:  498,  mentum;  499,  hypopharynx;  500,  antenna;  501,  mandible;  502,  spiracular 
disk 

Pupa:  503,  lateral  aspect;  504,  tip  of  pronotal  breathing  horn;  505,  male  cauda,  dorsal 
aspect 


1122 


MEMOIR  38 


PLATE  XCII 


TYPES    OF   ANAL   GILLS   IN   GENUS   TIPULA,    VENTRAL   ASPECT 

506,  Tipula  oropezoides;  507,  T.  nobilis;  508,  T.  caloptera;  509,  T.  dejecta-,  510,  T.  usitata; 
511,  T.  ignobilis;  512,  T.  iroquois  (supposition) 


1123 


22 
In 


1124 


MEMOIR  38 


523 


522 


PLATE 


xc* 

ft 


TIPULA    COLLABIS 


Larva:  519,  jnentum;  520,  hypopharynx;  521,  fifth  abdominal  segment,  dorsal  aspect; 
522,  spiracular  disk 

Pupa:  523,  male,  lateral  aspect 

1125 


MEMOIR  38 


PLATE  XCV 


528 


529 


527 


524 


525 

TIPULA   DEJECTA,    T.    USITATA,    AND    T.    TRIVITTATA 

Tipula  dejecta,  larva:  524,  men  turn;  525,  hypopharynx;  526,  spiracular  disk 

Tipula  usitata,  larva:  527,  fifth  abdominal  segment,  dorsal  aspect;  528,  spiracular  disk 

Tipula  trivittata,  larva:  529,  spiracular  disk 


1126 


1127 


4 


1128 


INDEX 

(When  more  than  one  page  reference  is  given,  the  more  important  references  are  italicized) 


A  PAGE 

Abdomen  of  pupa 756 

Abdominal  spiracles 757 

Adelphomyaria 895 

Adelphomyia 895 

americana 898 

cayui?a 898 

minuta 896 

Adipose  tissue 763 

Aeshnasoma 993 

riyertonensis 978,  993 

Amphibia  feeding  on  crane-flies 726 

Anal  gills  of  larva 748 

Antennae  of  larva 751 

Anthomyiidae  feeding  on  crane-flies 728,  732 

Antocha 799 

monticola 800 

saxicola 798,  800 

Antocharia 799 

Ants  feeding  on  crane-flies 729 

Arachnida  feeding  on  crane-flies 727 

Arachnida  parasitic  on  crane-flies ' 732 

Arrangement  of  families  and  lesser  groups .  .  .  701 

Asilidae  feeding  on  crane-flie* 728 

Aves  feeding  on  crane-flies 721 

B 

Bacteria  in  crane-fly  larvae 734 

Beling,  Theodor,  life  of 693 

Bibliography 1019 

Birds  feeding  on  crane-flies 721 

Bittacomorpha 783 

clavipes 784 

Bittacomorphella 779 

jonesi 780 

Bittacomorphinae 773,  779 

Bittacomorphinae,  keys  to  genera  of 779 

Body  form  of  larva 745 

Brachypremna 984 

dispellens 984,  985 

eocenica 984 

Breathing  horns  of  pupa 754 


Cheilotrichia  imbuta 

Chio'nea 

alpina 

araneoides 911, 

valga 

Circulatory  system 

Cladura 

delicatula 

flavof  erruginea 

Coleoptera  feeding  on  crane-flies 

Collecting 

Comparative  morphology 

Ctenophora. 

angustipennis 978, 

apicata 978, 

f  estiva 978, 

flaveolata 978, 

nigricoxa 978, 

pectinicornis 978, 

vittata 

Ctenophoraria 

Cylindrotoma 

distinctissima 960, 

splendens 705,960, 

tarsalis 


910 
950 
951 
951 
950 
760 
947 

.947,  948 
.   948 
....  729 
.  740 
....   766 
986 
986 


986 


967 
967 
967 


Cylindrotominae .• 

Cylindrotominae,  keys  to  genera  of . . 


PAGE 

792,     959 


Dactylolabaria  ........................... 

Dactylolabis  ............................. 

cubitalis  ....................  ......... 

denticulata  .................  .  .....  837, 

montana  .....................  ....:.. 

wodzickii  ........................  837, 

Dicranomyia  ........................... 

badia  ............................  820, 

dumetorum  ..................  798,  799, 

f  oliocuniculator  ...................  799, 

macateei  ............................ 

pilipennis  ................  .  ........... 

rara  ................................ 

simulans  ...........  .  .  ............  799, 

stulta  ...........................  820, 

trinotata  ........................  798, 

umbrata  ..................  '  .......  799, 

Dicranophragma  ......................... 

Dicranoptycha  ........................... 

winnemana  ......................  799, 

Dicranoptycharia  ...  ...............  ....... 

Dicranota  .....................  •  .......  899, 

bimaculata  .......................  895, 

Dicranotae  .............................. 

Dictenidia  ............................... 

bimaculata  .......................  978, 

Digestive  system  ......................... 

Diotrepha  ...............  ................ 

Diptera  feeding  on  crane-flies  .............. 

Djpterous  parasites  of  crane-flies.  ...  ....... 

Discobola  .......................  ........ 


caesarea  .........................  'yo, 

Ditomyiidae  ................  ............. 

•Dolichopeza  ............................. 

albipes  ..........................  978, 

americana  ........................... 

sylvicola  ....................  •  •  ...... 

Dolichopezaria  ........................... 

Duration  of  pupal  existence  ............... 


Economic  importance. 

Egg 

Elephantomyaria .... 

Elephantomyia 

westwoodi 


E 


. nr\o    ot\£i 

missa 798,  SOb, 

Ellipteraria 

Emergence  of  adult 

Empedomorpha 

empedoides •  ; 

Empididae  feeding  on  crane-flies 

Enemies  of  crane-flies 

Epipharynx  of  larva 

Epiphragma . •  •  •  •  • • • • 

fascipennis <*>'•  ?**• 

picta 837' 

solatrix 

Epiphragmaria 

Eriocera w^V 

cinerea xxi' 

fultonensis tMO> 


852 
852 
854 
853 
855 
854 
819 
824 
819 
820 
820 
820 
820 
820 
822 
819 


828 
906 
906 
906 
987 
987 
758 
800 
727 
731 
815 
815 
815 
788 
981 
981 
982 
978 
981 
720 


736 
714 
952 
952 
953 
806 
807 
806 
710 


721 
750 
843 
846 
844 
844 
843 
881 
886 
890 


1129 


1130 


CHARLES  PAUL  ALEXANDER 


Eriocera  (continued) 

longicornis 

spirosa 

Eriocera,  keys  to  species  of 

Erioptera 

armata 

caloptera 

chlorophylla 


PAGE 

.  .  704,  837,  888 
837,  883 

882 

914 
921 
922 
918 
915 
915 
922 
915 
914 
915 
922 
920 
915 
919 


PAGE 

Hexatomaria,  keys  to  genera  of 877 

Hexatomini 835 

Hexatomini,  keys  to  subtribes  of 836 

Histoblasts 763 


915, 
910, 
910, 


................  915, 

................  915, 

.................  915, 

................  910, 

graphica 

knabi 

lutea 

maculata 

megophthalma 

parva  ............................... 

septemtrionis  .....................  915, 

sordida  ..........................  910, 

vespertina  .......................  915, 

Erioptera,  key  to  pupae  of  .................      915 

Eriopteraria  .............................     911 

Eriopterine  No.  1  ........................      956 

Eriopterini  ..............................      908 

Eriopterini,  keys  to  genera  of  ..............      909 

Eulimnophila  ............................      872 

External  morphology  of  larva  ..............      744 

External  morphology  of  pupa  ..............      752 

External  parasites  ........................      730 


Fat  bodies,  or  adipose  tissue 763 

Feeding  habits  of  adults 713 

Feeding  habits  of  larvae 717 

Fish  feeding  on  crane-flies 726 

Flagellate  parasites  of  crane-flies 734 

Fungous  parasites  of  crane-flies 732 


G 

General  features  of  larva 744 

General  features  of  pupa 752 

Geological  record 764 

Geranomyia 816 

canadensis 817 

diversa 817 

rostrata 817 

virescens 817 

Gnophomyia 934 

rufa 911,  934 

tripudians 911,  934 

tristissima 911,  935 

Gonomyella 938 

Gonomyia 938,  940 

alexanderi 938,  939 

kansensis 938,  941 

subcinerea 938 

sulphurella. 938,  940 

tenella 911,  938 

Gonomyia,  key  to  pupae  of 938 

H 

Habromastix  cinerascens 980 

Head  of  larva 748 

Head  of  pupa 753 

Helobia 928 

hybrida 911,  929 

macroptera 929 

Hexatoma 877 

bicolor 878 

megacera 838,  878 

nigra 838,  878 

saxonum 878 

Hexatomaria 876 


Historical  summary  of  immature  stages .  . 

Holorusia 

rubiginosa 978 

Hoplolabis 

Hymenoptera  feeding  on  crane-flies 


735 
993 


921 
729 


Imaginal  disks 763 

Internal  morphology 758 

Internal  parasites  of  crane-flies 733 


Killing  and  preserving  specimens 740 


Labium  of  larva 750 

Labrum  of  larva 750 

Larva 715 

Larval  habitat 716 

Lasiomastix 863 

Leiponeura 939 

Libnotes 814 

perkinsi 798,  814 

Life  activities  of  crane-flies 710 

Life  histories  of  crane-flies,  representative .  .  .  704 

Limnobaria 808 

Limnobaria,  keys  to  genera  of 808 

Limnobia 809 

annulus 798,  809 

bifasciata 798,  809 

cinctipes 810 

decemmaculata 798,  809 

fallax 810,  813 

.    flavipes 798,  810 

immatura 798,  810 

indigena 810 

inusta 798 

macrostigma 798,  810 

nigropunctata 798,  810 

nubeculosa 798,  810 

obscuricornis 798,  810 

parietina 810 

quadrimaculata 798,  809 

sexpunctata 798,  810 

solitaria 810 

triocellata 798,  810,  814 

tripunctata 798,  810 

xanthoptera 798,  809 

Limnobiinae 793 

Limnobiinae,  keys  to  tribes  and  subtribes  of .  793 

Limnobiini 795 

Limnobiini,  keys  to  subtribes  of 797 

Limnophila 858,  868 

adusta 867 

bryobia 837,  869 

dispar 837,  860 

f erruginea 837,  860,  866 

fuscovaria 861 

hyalipennis 837,  860 

lineola 837,  860 

macrocera 864 

nemoralis 837,  860 

ochracea 837,  860 

pallida 837,  860 

pictipennis 837,  869 

punctata 837,  868 

sinistra 837,  859 


THE  CRANE-FLIES  OF  NEW  YORK  —  PART  II 


1131 


Limnophila  (continued)  PAGE 

ultima 711 

unica 860 

Limnophilae 858 

Limnophilaria 856 

Limnophilaria,  keys  to  genera  and  subgenera  of  857 

Liogma 960,  969 

glabrata 960,  969 

nodicornis 961,  971 

Liogma,  keys  to  species  of 970 

Lipsothrix  icterica " 910 

Longurio 990 

testaceus 991 

M 

Mammalia  feeding  on  crane-flies 721 

Mandible  of  larva 751 

Mating 710 

Maxillae  of  larva 752 

Mesocyphona 921 

Methods  of  rearing 741 

Methods  of  study 741 

Molophilus 911 

bifilatus 910,  912 

hirtipennis .  .  912 

obscurus 910,  911 

ochraceus 910,  911 

ursinus 914 

Mongoma 944 

Muscular  system 763 

Mycetobia 788 

Mycetobiinae 788 

N 

Neanotype 743 

Neocladura 947 

Nephrotoma 1016 

analis 980,  1017 

cornicina 980,  1017 

crocata 980,  1017 

eucera 1017 

ferruginea 737,  981,  1017 

histrio 980 

iridicolor 980 

lineata 980,  1017 

lunulicornis 980,  1017 

maculata 980,  1017 

polymera 1017 

pratensis 980,  1017 

quadrifaria 981,  1017 

virescens 1017 

Nepionotype 743 

Nervous  system 762 


Odonata  feeding  on  crane-flies 727 

Orimarga 799 

Orimargula 799 

Ormosia 922 

haemorrhoidalis 910,  923 

innocens 923,  925 

lineata 

meigenii 

nigripila 

nodulosa 910,  923 

nubila 923 

varia 910,  923 

Ormosia,  key  to  pupae  of 923 

Or  jpeza 982 

obscura 983 

Oviposition 713 


Paramqngoma 

Parasitic  natural  enemies  of  crane-flies    . 

Pedicaria 

Pedicaria,  key  to  genera  of ... 

Pedicia 

albivitta . .895, 

cpntermina 

rivosa 895, 

Pediciae 

Pediciini 

Pediciini,  key  to  subtribes  of 

Penthopter'a 

albitarsis 838, 

Phalacrocera 

replicata 960, 

tipulina 

Phylidorea 

Phylogenetic  considerations 

Phylogeny 

Pilaria 

discicollis 837, 

fuscipennis 837, 

quadrata 

recondita 

tenuipes 837, 

Pilaria,  keys  to  species  of 

Pisces  feeding  on  crane-flies 

Polymera 836, 

geniculata 

georgiae 

magnifica 

Poly  meraria 836, 

Predatory  natural  enemies  of  crane-flies 

Prionocera 

fuscipennis 978, 

parri 978, 

Progonomyia 

Protoplasa 

«    fitchii 

Pseudolimnophila 

inornata 

luteipennis 837, 

Pseudolimnophilaria 

Ptychoptera 

albimana 773, 

contaminata 

lacustris 

lenis 773, 

paludosa 

rufocincta 

Pty  chopteridae 

Ptychopteridae,  keys  to  subfamilies  of 

Ptychopterinae 

Pupa 

Pupal  duration 


PAGE 

944 
730 


900 
900 
900 


894 


961 

962 
961 
866 
766 
763 
872 
872 
872 
875 
874 
873 
872 
726 
893 
893 


721 
995 


938 
769 
769 
848 
851 
8JO 
848 
773 
774 
773 
773 
775 
773 
775 
772 
773 
773 
719 
720 


..910,     923 

923,     928 

923,     927 


References  cited 

Reproductive  system 

Respiratory  organs 

Resting  habits 

Rhabdomastix 

flava 

schistacea 911, 

Rhamphidaria 

Rhamphidia 

flavipee 799, 

longirostris 799, 

mainensis 

Rhaphidolabina 

flaveola 


1019 
762 
760 
712 
942 
942 
942 
830 
830 
833 
830 
831 
901 
901 


1132 


CHARLES  PAUL  ALEXANDER 


Rhabidolabis  

PAGE 

907 

Tipula  (continued) 
lunata  

PAGE 

979,     999 

cayuga  

907 

luteipennis  

979,     999 

tenuipes  

895,     907 

lutescens  

978 

Rhipidia  

825 

macrolabis  

1014 

bryanti  

826 

marmorata  

979,     999 

domestica  

799,     886 

maxima  

979,     999 

fidelis  

826 

micans  

979 

macula  ta  
uniseriata  

799,     826 
799,     826 

nigra  
nobilis  

.  737,  979,     999 
1004 

Rhyphidae  

787 

nubeculosa  

979,     999 

Rhyphinae  

787 

ochracea  

979,     999 

Rhyphus  

787 

oleracea  

.  737,  979,     999 

punctatus  

788 

oropezoideo  

1001 

pabulina  

979,     999 

s 

pagana  

979,     999 

Sacandaga  
Scatophagidae  feeding  on  crane-flic 
Spiracular  disk  of  larva  
Sporozoan  parasites  of  crane-flies.  . 

.      942 
s  .  ..     728 
.....      746 
734 

paludosa  
parva  
peliostigma  
pruinosa  

.  737,  979,     999 
737,     979 
979,     999 
979,     999 

Styringomyia  

957 

rufina  

.710,979,     999 

ceylonica  

958 

sayi  

1000 

didyma  

958 

scripta  

.-  979,     999 

jacobsoni  

958 

selene  

.979,  999,  1016 

venusta  

958 

serta  

980 

vittata  

958 

signata  

979      999 

Styringomyiini  

957 

simplex  

738 

Swarming  

711 

subnodicorais  

979,     999 

taughannock  

1013 

T 

tephrocephala  

1000 

tricolor  

1000 

Tachinid  parasites  of  crane-flies  .  .  . 
Tanyderidae  

731 
.      769 

trivittata  
truncorum  

..980,  1009 
979      999 

Tanyptera  

988 

ultima  

980,  1001 

atrata  

978,     988 

umbrosa  

1000 

atrata  ruficornis  

978,     988 

unca  

980,     999 

flavicornis  

978,     988 

usitata  

1008 

f  rontalis  

988 

varncorms  

980,     999 

f  umipennis  
Teucholabis  

978,     988 
945 

varupennis  
vernalis  

980,     999 
980,     999 

complex*  

911,     946 

vittata  

980,     999 

Thaumastoptera  

716,     798 

winnertzii  

980 

Thorax  of  pupa  
Tipula...  ......  f  

754 
998 

Tipularia  
Tipulidae  

990 
791 

abdominalis  

980,  1011 

Tipulidae,  keys  to  subfamlliei  of  .  . 

792 

angustipennis  

1000 

Tipulmae.  .  

792,     974 

annulicornis  

980 

Tipulinae,  keys  to  genera  of  

977 

arctica  

980,  1001 

Tipulini  

.     981 

bella  

1004 

Tipulodina  pedata  

978 

bicornis  

.  738,  980,  1001 

Tipuloidea,  keys  to  families  of  .  .  .  . 

.........      768 

caesia  _  

978,     999 

Toxorhina  

.     955 

caloptera  

980,  1005 

madagascariensis  

955 

cayuga  

1000 

muliebris  

956 

collaris  

1002 

Trentepohlia  

943 

cunctans  

.  738,  980,  1001 

bromeliadicola  

911,     944 

dejecta  

1006 

crucif  erella  

943 

dilatata  

978,     999 

humeralis  

943 

eluta  

980,  1001 

leucoxena  

911,     945 

flavicans  
flavolineata  

980 
978,     999 

pennipes  
Trichocera  

911,     944 
789 

f  uliginosa  

1000 

'f  uscata  

790 

fulvipennis  

978,     999 

hiemalis  

.      790 

gigantea  

979 

regelationis  

790 

hortensis  

.....978,     999 

Trichocerinae  

788,     789 

hortulana  

979,     999 

Trichotipula  

1001 

ignobilis  

1010 

Tricyphona  

903 

imbecilla  

979 

auripennis  

.      904 

infuscata  

980 

immaculate  

895,     904 

iroquois  

1014 

inconstans  

904 

irrorata  

979,     999 

paludicola  

.     903 

lateralis  

979,     999 

schineri  

895,     904 

longicornis  

980 

vernalis  

904 

THE  CRANE-FLIES  OF  NEW  YORK — PART  II 


1133 


PAGE 

Trimicra 932 

pilipes 911,  Q32 

reciproca 933 

Triogma 973 

exculpta 974 

kuwanai 974 

trisulcata 961,  974 

Types  of  the  immature  stages 743 


Ula. 


U 


Ula  (continued)  PAGE 

elegans 837,  840 

macroptera 837,  839 

Ularia 838 

Ulomorpha 869 

pilosella 869 

Ulomorphae 869 


W 


boLtophlla . 


837,     840        Wasps  feeding  on  crane-flies. 


729 


GENERAL  LIBRARY 
UNIVERSITY  OF  CALIFORNIA— BERKELEY 

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Renewed  books  are  subject  to  immediate  recall. 


SEP     8  1954 

HOV281952 


LD  21-100m-l,'54(1887sl6-)476 


